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BuLtouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

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FiscHEer, P. H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

Koun, A. J. 1960@. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

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Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 104 Band

June 1994 Junie

Part 2 Deel

PHYLLODACTYLUS PERINGUEYI

(REPTILIA, GEKKONIDAE)

ITS TAXONOMIC HISTORY, REDISCOVERY,

AND PHYLOGENETIC AFFINITIES

WILLIAM R. BRANCH

AARON M. BAUER

Cape Town Kaapstad

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D3103

PHYLLODACTYLUS PERINGUEYI (REPTILIA, GEKKONIDAE)

ITS TAXONOMIC HISTORY, REDISCOVERY,

AND PHYLOGENETIC AFFINITIES

WILLIAM R. BRANCH

Port Elizabeth Museum, P.O. Box 13147, Humewood, South Africa 6013

AARON M. BAUER

Department of Biology, Villanova University, Villanova, Penn. 19085, USA

(With 5 figures and 1 table)

[MS accepted 20 August 1993]

ABSTRACT

The taxonomic status and provenance of the two syntypes of Phyllodactylus peringueyi

are reviewed. A redescription of the species is given, based on the extant syntypes and new

material referable to the species. The species is compared with all known geckos having

Phyllodactylus-type toes. It is not referable to any other known species group and, on

biogeographic grounds, it seems likely that P. peringueyi forms part of an endemic southern

African radiation of leaf-toed geckos.

A lectotype and paralectotype are designated from the extant syntypes, which survive in

poor condition. Of the two localities documented for the syntypes, i.e. Namaqualand and

Chelsea Point, the first is considered doubtful. Many other specimens accessioned at the same

time, and purportedly also collected by Péringuey from Namaqualand, represent species not

known from that region and were probably inaccurately documented. There is no indication

that the Chelsea Point locality is inaccurate, although no additional specimens have been

collected from the region despite intensive searches. However, the species has recently been

rediscovered 77 km west of Chelsea Point, in salt marshes of the Kromme River estuary,

Eastern Cape.

CONTENTS

PAGE

In froductiOnie aa cacsscs cr sac etme neo celacecapiowcaacmacindec eee ciee tans ene sae 14

MaterialvandsmethodSis2scc-csccasscen ccc doc cee vec sone ceaealeccuiseece sutee chic aaetas 14

FRESuIts' and GISCUSSION I acess chc cnn cena necessities eae Sone oiauisis sitelaree ete sincrataats 15

Condition of the syntypes of Phyllodactylus peringueyi ................++. 15

Designation of lectotype and paralectotype ).2 2 -..teeee case sece-seeces 17

REdISCOVEnyAOLtHe SPECIES) een sete ceases ee stian: seitemitsmr saints cesar a

Redescription Of, the Species. en ce.c sees esceetenccneh clin cciiseinsccoes ncaa te 19

History, of-the’syntypes and their provenance 2.0. 6.0.0.2. cccueeewceooenee 22

The origin of the syntypes of Phyllodactylus peringueyi ...............++. 23

The distribution and habitat of Phyllodactylus peringueyl ..............+. 25

Relationships of Phyllodactylus Peringueyi .........0.cecceceecees ene eeeees PLY]

ACKNO WIERD SEMENES Hic sociisiseeeee accu cne eee oes e ewer ete brancclneaaaeseinn 28

FRELEL ENCES pees ti a eeiaaatceperann cater Muietelon slace eine ato netieiad SESE sce aeea sae sigeta sat 28

PRET ixars ore sece cries sean cam See cac oem cee seis sree aioe Sense bidoe Merce toss ettied 30

Ann. §S. Afr. Mus. 104 (2), 1994: 13-30, 5 figs, 1 table.

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 14

INTRODUCTION

The genus Phyllodactylus is a widespread and probably polyphyletic assem-

blage of small to moderate-sized geckos, with representatives world-wide.

Recently, certain monophyletic groups have been recognized within Phyllo-

dactylus sensu lato and have been accorded generic status (Dixon & Anderson

1973; Dixon & Kroll 1974; Kluge 1983; Cogger 1986). Species currently

retained in Phyllodactylus are chiefly distributed throughout the New World,

with the greatest diversity in the arid regions of the Pacific coast of Mexico and

South America, and in Venezuela. The remaining Old World Phyllodactylus

include two species in Thailand, one in the Mediterranean region, two on the

island of Socotra, and five described species in southern Africa. Phyllodactylus

ansorgii, endemic to Angola, is relatively poorly known but two of the South

African species, P. lineatus and P. porphyreus, have been well collected and

their distributions are well established, and a third, P. microlepidotus, is

becoming so.

Since its description at the beginning of this century, the status and distribu-

tion of Phyllodactylus peringueyi Boulenger, 1910, have remained a persistent

problem in South African herpetology. The two syntypes, which are believed to

have been acquired 19 years apart, are catalogued as coming from two widely

separated localities (Little Namaqualand and the Eastern Cape). Few southern

African lizards have comparable distributions and these are common, wide-

spread generalists with wide habitat tolerances (e.g. Mabuya capensis and

Agama atra). No known geological or climatic events in the history of southern

Africa are likely to have generated relict distributions matching the stated

localities of the syntypes. In addition, the morphology of P. peringueyi is unlike

that of any other South African gecko currently assigned to Phyllodactylus.

Prior to this report, no new specimens had been discovered since the type

description.

Although several authors have reviewed the problem posed by these enig-

matic specimens (Hewitt 1937; FitzSimons 1943; McLachlan 1988), no new

information regarding their identity has been available and, to date, no attempt

to use any but the most superficial aspects of pholidosis in determining their

affinities has been made. We present here the results of our studies on: the prov-

enance and taxonomic history of the species; the condition and a fuller descrip-

tion of the syntype specimens; morphological variation in recently collected

fresh material referable to the species; and a preliminary comparison of the

species with all known congeners and species from allied leaf-toed genera.

MATERIALS AND METHODS

External features were examined using a binocular dissecting microscope,

and osteological features revealed through the use of X-rays. Measurements

were taken with Brown and Sharpe ‘Digit-Cal Plus’ digital calipers.

The syntypes of Phyllodactylus peringueyi and all fresh material referable to

this species were examined (see Appendix). In addition, all known species of

gekkonine geckos possessing ‘Phyllodactylus-type’ toes (Russell & Bauer 1989)

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 15

were compared externally to address the specific distinctness of P. peringueyi.

The species lists of Kluge (1983, 1991) served as a guide to the recognized taxa.

Specimens of nearly all members of the genera Paroedura, Asaccus, Ebenavia,

Christinus and Urocotyledon, in addition to those currently assigned to Phyllo-

dactylus sensu stricto, were examined in the following collections: British

Museum (Natural History) (BMNH), California Academy of Sciences (CAS),

University of Kansas Museum of Natural History (KU), Academy of Natural

Sciences of Philadelphia (ANSP), Louisiana State University Museum of

Natural Sciences (LSU), United States National Museum (USNM), Field

Museum of Natural History (FMNH), Museum fiir Naturkunde der Humboldt-

Universitat zu Berlin (ZMB), Port Elizabeth Museum (PEM), and South

African Museum (SAM). Specimens of the following species were not

examined, but were compared on the basis of published descriptions: Phyllo-

dactylus bordai, P. rutteni, P. transversalis, Paroedura homolorhinus, P. ovi-

ceps, and Urocotyledon weileri. In addition, osteological comparisons were

made using cleared-and-stained and dry skeletal material from the afore-

mentioned institutions, as well as X-rays.

Historical information regarding the syntypes of Phyllodactylus peringueyi

was derived from the surviving correspondence, registers, annual reports, and

catalogues in the South African Museum, covering the period 1885-1910,

during which time the specimens were acquired and described.

RESULTS AND DISCUSSION

Phyllodactylus peringueyi Boulenger, 1910

CONDITION OF THE SYNTYPES OF PHYLLODACTYLUS PERINGUEYI

The type description (Boulenger 1910) is based on two specimens, both of

which are extant and housed in the herpetological collection of the South

African Museum, Cape Town. They are now in only poor condition. Both were

initially fixed mn contorted positions, and numerous subsequent attempts to study

their anatomy have resulted in considerable damage. This includes both

incidental damage during dissections to determine internal features, and

accidental damage in handling and during transit.

SAM-777. Snout-vent length, 23.5 mm; mid-body diameter, 4.5 mm. The

skin is torn on the throat and in the middle of the belly, and has been cut around

the margin of the lower jaw. The lower portion of the left forelimb, distal to the

elbow, is absent (Fig. 1A). Similarly, only a tibia remains of the lower portion

of the right hind limb. The right fore-limb is broken and torn at the elbow,

although the lower part of the limb remains attached by a small skin flap. The

specimen has no tail, this being lost distal to the hémipeneal pouches. The

description of FitzSimons (1943) contains no comment on the tail. However, in

the original description, Boulenger (1910) noted a total length of 45 mm (tail

regenerated), but did not state which specimen was measured. A detached tail

(11 mm) was found in the bottle containing the syntypes; it is likely to have

come from SAM-777.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-8628. Snout-vent length, 28.1 mm. Damage to this specimen is more

extensive. Most of the tail is missing, leaving only the proximal 3 mm. The

right fore-limb is broken just distal to mid-tibia, although the distal portion is

attached by a skin flap (Fig. 1B). The thumb and terminal phalanges of the

fourth finger are missing. The right hind limb is also broken just below the

knee, with the lower portion of the limb attached by a skin flap. Only the third

and fourth digits remain on the left fore-limb, and the middle digit of the left

hind limb is truncated at the second phalanx. The skin over much of the dorsal

surface of the skull, from the tip of the snout to the middle of the parietal

region, has been removed. The right eyeball is punctured and the skin around

the right margin of the lower jaw has been cut.

Fig. 1. Radiographs (dorsal views) of the syntypes of Phyllodactylus peringueyi.

A. Lectotype SAM-777. B. Paralectotype SAM-8628. Note the reduced number of

presacral vertebrae. The arrow in A indicates the cloacal bone in the male. The dark

image of the hind limb is a result of over-exposure of the X-ray positive print to

bring out details of the digits. Actual size of specimens: A. 23.5 mm SVL;

B. 28.1 mm SVL.

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI NG

DESIGNATION OF LECTOTYPE AND PARALECTOTYPE

Lectotype. We here designate the adult male, SAM-777, stated to have

been collected by L. Péringuey in Little Namaqualand, Cape Province, South

Africa, as the lectotype of the species.

Our reasons for this are as follows: it is in better condition, retaining large

areas of skin covering the head; it is a male and thus displays a number of

potentially useful sexually dimorphic features (e.g. cloacal bones); the detached

tail most likely came from SAM-777 and this, therefore, is probably the

specimen that formed the basis for measurements listed in the type description.

Finally, although Boulenger (1910) did not designate a holotype, and none has

been subsequently designated, the catalogue entry for SAM-777 has a pencilled

entry ‘type’.

Paralectotype. SAM-8628, an adult female, reputedly collected by

A. Moorhouse at Chelsea Point, near Port Elizabeth, Cape Province, South

Africa. FitzSimons (1943) was mistaken in describing SAM-8628 as a male.

Although we consider the stated locality for the lectotype to be mistaken and

that for the paralectotype to be more acceptable, for reasons discussed below

(see p. 23), we caution against acceptance of the published localities of both

syntypes. The species appears to be restricted to the Eastern Cape (see p. 25).

REDISCOVERY OF THE SPECIES

Prior to this report, no new specimens of P. peringueyi have been collected

since the original type description (Boulenger 1910). Due to its problematic

status, the species was listed as ‘Indeterminate’ in the South African Red Data

Book—Reptiles and Amphibians (McLachlan 1978; revised Branch 1988c), and

even described as ‘one of the herpetological mysteries of the region’ in the most

recent synopsis of the subcontinent’s reptilian fauna (Branch 1988a: 211).

Recently, whilst this manuscript was in preparation and after a preliminary

synopsis of its findings had been presented (Branch & Bauer 1992), three speci-

mens of a small, striped, leaf-toed gecko with keeled dorsal tubercles (Fig. 2),

were collected in transitional salt-marsh/floodplain vegetation (including Sarco-

cornia perennis, Chenolea diffusa and Iporobolus virginicus) along the banks of

the lower Kromme River (3424Bb Humansdorp; 34°07'15"S 24°45'30"E; alt.

1 m asl.). The specimens were referable to P. peringueyi. Fuller details of the

species’ rediscovery have been published elsewhere (Branch et al. 1992).

Subsequent trips to the region have shown the species to be common along the

banks of the lower Kromme River (see Fig. 5), but it has not been found in

adjacent estuaries. It shelters under logs or in tussocks of thick vegetation,

particularly Restio sp. In habitus, colour and preferred habitat of tussock

vegetation, it shows a striking similarity to some Australian diplodactyline

geckos, particularly Crenadactylus ocellatus and species of the Diplodactylus

michaelseni group. There are also intriguing similarities with the small

Madagascan gecko Ebenavia inunguis. An analysis of the species’ distribution

and preferred habitat is in preparation (Branch in prep.).

ANNALS OF THE SOUTH AFRICAN MUSEUM

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REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 9

REDESCRIPTION OF THE SPECIES

FitzSimons (1943) provided a description of the species, modified from that

of Boulenger (1910). We have been able to determine a number of additional

features that either extend or correct previous descriptions.

The head is flattened above; the snout is elongate and pointed, and almost

twice the diameter of the eye and a little longer than the distance between the

eye and the ear-opening (Fig. 3). The latter is small and rounded, whereas the

eye has a vertical pupil. Head measurements of the syntypes are: length of head

(tip of snout to rear of skull), 6.9 mm (SAM-777) and 7.3 mm (SAM-8628);

width of head (at rear), 4.2 mm (both syntypes). Boulenger (1910) and Fitz-

Simons (1943) described the head as being 1.5 times as long as broad, but the

syntype measurements give a slightly higher ratio (1.65 and 1.73). Fresh

material gives an even higher ratio (males, n = 5, range 1.78-1.97, mean 1.87;

females, n = 4, range 1.78-1.93, mean 1.85), with no evidence of sexual

dimorphism.

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& Sa

“C a ese CLES RSS

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Seat a

Fig. 3. Profile of Phyllodactylus peringueyi showing details of scalation

(based on PEM R6908; lower Kromme River, Humansdorp District,

Eastern Cape).

Boulenger (1910) and FitzSimons (1943) described the nostril as pierced

between the rostral and four small nasal scales. However, in all fresh material,

the nostril pierces a small nasal scale that is bordered by the rostral, first upper

labial, two supranasal (the anterior larger and slightly wider than long), and by

2-3 small postnasal granules. Damage to the snout region of both syntypes

makes it impossible to determine whether the original type description of the

nasal condition is correct. The narrow borders of the nasal scale that surrounds

the nostril make it easily overlooked.

The rostral is twice as broad as deep and lacks a median cleft. Boulenger

(1910) noted that the granules on the snout were keeled, and FitzSimons (1943)

added that they were larger and more strongly keeled than the scales on the

back of the head. However, in fresh material the snout granules vary in the

extent of their keeling and are equal to, or only slightly larger than, the scales

on the crown of the head. The latter are also more irregular in outline. Other

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

details of head scalation include: 13-14 scales around ear; usually 13 scales

between anterior margin of ear and posterior margin of eye; 9-11 scales

between nostril and anterior margin of eye; 2 scales between upper labials and

eye; 21-24 scales (excluding circumorbitals) between eyes across crown of

head; 6-7 upper labials; 6-8 lower labials; no enlarged chin shields; 2-3 scales

bordering the mental; 2-3 granules bordering rostral between anterior

supranasals.

The body is rather elongate; the back is covered with coarse, more-or-less

keeled granules, intermixed with numerous enlarged, longitudinally oval,

strongly keeled tubercles. These are scattered, although they may form stag-

gered irregular rows on the upper flanks. There are 13-14 tubercles across the

back (11-12 in SAM-8628, which are also less prominently keeled). The gulars

are small, rounded and granular, whereas the belly scales are smooth, flat, cyc-

loid, imbricate, and at least twice the size of the gulars. The scales beneath the

proximal portion of the tail are almost twice the size of those on the belly, and

16-18 scales border the anterior margin of the cloaca. There are no preanal

pores. A large, semicircular scale, almost twice the size of adjacent scales, pro-

trudes on each side of the tail base, just distal to the cloaca, forming a cloacal

spur. It is present in both sexes, but is larger and more protruding in the male

lectotype and other sexually mature males. The latter also have a distinctly

swollen tail base due to the presence of the hemipenes.

Fig. 4. Underside of right pes of Phyllodactylus peringueyi

showing details of scalation (based on PEM R6886; Kaila de

Balaia, Kromme River, Humansdorp District, Eastern Cape).

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 21

The limbs are short and pentadactyl (Fig. 4). The distal scansors are

rounded, moderately large, and medially divided, with a small claw between the

paired distal expansions of each toe. The median row of subdigital scales is well

developed and transversely enlarged, with 6-7 scales beneath the longest digit

of the pes.

Colour

Superficially, there may appear to be sexual dichromatism; seven of eight

adult males were boldly striped, whereas most females (five of seven) and all

juveniles (two) and hatchlings (two) were uniformly coloured. However, captive

specimens of both sexes readily change the intensity of the striped pattern, and

gravid females may darken considerably. The correlation between coloration

and activity and/or reproductive condition needs to be investigated further.

Striped phase (based on PEM R6886, an adult male). Body above with a

light golden-brown vertebral stripe (1.4 mm wide, including 2-3 rows of dorsal

tubercles with intervening granules) that extends from the back of the head to

the beginning of the regenerated tail (in specimens with original tails, the stripe

extends along the top of the tail for its full length). On each side this is edged by

a thin black band (2 granules wide), which in turn is bordered by a broad

(3-5 keeled tubercles wide) dark-brown dorsolateral band. Anteriorly, the latter

is faintly divided by a very thin (1-2 granules wide) light-brown, longitudinal

stripe that extends to the nostril, whereas posteriorly the band breaks up into

fine, anastomosing stripes on the lower flanks. The dorsolateral bands fuse on

the crown of the head at the level of the eyes; posteriorly they widen consider-

ably and are most prominent over the hind limbs and tail base. The flanks are

covered with a pale lateral ‘band’ that is composed of 3-4 faint, thin (one gran-

ule wide) brown stripes, separated by tan lines (also one granule wide) that

occasionally anastomose. The band extends as a golden-brown stripe through

the temporal region and eye, to the nostril. In some males the vertebral stripe

lacks the black edge.

The iris is golden, coppery red, whereas the upper and lower labials are

dirty cream, heavily stippled with dark spots. The throat has 4-6 vague, thin

. (one granule wide), dark-brown longitudinal stripes. The belly is also dirty

cream with vestiges of thin dark stripes that radiate on to the lower surfaces of

the hind limbs. The upper surfaces of both fore- and hind limbs are dirty golden

brown, heavily stippled with dark spots.

Uniform phase (based on PEM R6908, an adult female). Head, body and

upper surfaces of the limbs and tail uniform brown. There are faint vestiges of

thin dark stripes on the flanks. The throat, belly and upper and lower labials are

dirty cream with faint dark speckling.

Dimensions

Maximum snout-vent length: 28.3 mm females, 27.8 mm males. Unregen-

erated tails are approximately equal in length to that of the snout-—vent;

regenerated tails are usually smaller. Hatchlings measure 13-14 mm SVL, with

a proportionately shorter tail (9-10 mm).

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

HISTORY OF THE SYNTYPES AND THEIR PROVENANCE

Cataloguing procedures and data recording in nineteenth-century herpeto-

logical collections were generally imprecise, and have generated several prob-

lems regarding type localities within Phyllodactylus (see Dixon & Huey 1970).

However, the special circumstances at the South African Museum (SAM) have

added exceptionally to the confusion surrounding Phyllodactylus peringueyi.

The lectotype, SAM-777, was acquired during the period that Péringuey

himself was Assistant Director at the SAM. According to Summers (1975: 95),

Péringuey had an ‘. . . incurable habit of trusting to his memory for the prov-

enance of finds of all sorts’, and once remarked ‘I don’t know what this

Museum would do without my memory’. SAM-777, along with many reptiles

supposedly collected in southern Namibia and the Western Cape, was not cata-

logued until 4-7 September, 1896, after the arrival of W. Sclater as Director at

the South African Museum. At that time a large series of specimens (SAM-733

to 826, 835 to 892), all bearing the locality ‘Namaqualand’ and collector

L. Péringuey, were accessioned.

The original entry in the herpetological catalogue, in the handwriting of Mr

R. M. Lightfoot, general clerk and bookkeeper at the SAM, reads:

‘SAM-777, Phyllodactylus porphyreus var. Namaqualand, 4 Sept 1886,

L. Péringuey, in alc.’

A later note, in unidentified handwriting, adds:

‘Now P. peringueyi Boulenger, type.’

‘In 1885 Annual Report’ has also been added in pencil by G. McLachlan,

Curator of Herpetology, SAM, 1976-1989.

McLachlan believed that SAM-777, one of the P. peringueyi syntypes, was

among the specimens listed in an appendix (in unknown handwriting) to the

1885 Annual Report of the SAM listing herpetological acquisitions for that year.

These include seven species with the Namaqualand locality:

‘Chondrodactylus angulifer, Phyllodactylus sp. nov. ?, Phyllodactylus sp.

nov. ?, Ptenopus garrulus, Pachydactylus formosus, Phyllodactylus lineatus

Gray, Pachydactylus bibronii and Agama aculeata.’

A register of specimens for 1885 (without localities or specimen numbers)

lists, in Péringuey’s handwriting:

‘1 Chondrodactylus angulifer, 2 Phyllodactylus lineatus Gray, 4 Pachydac-

tylus formosus Smith, 1 Phyllodactylus n. sp., 4 Pachydactylus bibronii,

1 Pachydactylus n. sp., 4 Ptenopus garrulus, Agama aculeata.’

The two lists differ only in that the report mentioned two specimens of a

new species of Phyllodactylus, whereas the register gives one new Phyllo-

dactylus and one new Pachydactylus. Whether the specimens were re-identified

or incorrectly transcribed into the Annual Report from Péringuey’s register is

unknown.

The paralectotype is accessioned in the SAM herpetological catalogue as:

‘SAM-8628 Diplodactylus sp. ? nov. Chelsea Point, nr. Port Elizabeth, 2 Dec

1904, A. C. Moorhouse, alc. Caught on beach, see letter 2/12/1904.’

Again, a later note adds ‘Now P. peringueyi Boulenger.’

The 1904 SAM Annual Report notes, under thanks for recent acquisitions:

‘Mr A. Moorhouse, for a new Lizard, of the family Geckonidae, and the genus

Diplodactylus, found by him near Port Elizabeth. This is a specially interesting

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 23

form, as all the other species of the genus hitherto known have been obtained in

Australia.’

Unfortunately, the letter from Moorhouse accompanying the specimen when

it was donated to the SAM is not on file (M. Cherry pers. comm.). However, a

letter from SAM Director W. Sclater to G. A. Boulenger at the British

Museum, undated but bound between letters dated 30/11/1904 and 7/12/1904 in

the SAM Letter Book for November 1897-October 1905, was located. In it

Sclater asked Boulenger to identify a number of problematic specimens (several

fish, a snake, and two geckos): ‘I am sending you a small tube of things I cannot

make (anything) of. The little Phyllodactylus from Algoa Bay seems quite differ-

ent from (anything) we have. It appears to me to be (near) tuberculosus of your

Catalogue . . . [SAM] 8628 Seashore on rocks of Algoa Bay, 12 miles from

Port Elizabeth.’

THE ORIGIN OF THE SYNTYPES OF PHYLLODACTYLUS PERINGUEYI

In the description of Phyllodactylus peringueyi, Boulenger (1910: 494) pro-

vided the following details regarding the origins of the two syntypes: ‘A male

from Little Namaqualand (coll. Péringuey) and female from Port Elizabeth

(coll. Moorhouse).’ Distributions of African reptiles were poorly understood at

the time, but it has since become clear that at least one of the localities is prob-

ably in error. Because Péringuey did not keep a written record of specimens

accessioned, it was not until 1896, as many as 12 years after their collection,

that specimens were catalogued. Many opportunities for mistakes in subsequent

documentation of specimens may have occurred. This casts doubt especially on

the ‘Namaqualand’ locality of the lectotype.

The term ‘Namaqualand’ was used by early workers in a much broader

sense than it is today. As understood at the time of Péringuey, it probably

included the area from Walvis Bay south to Clanwilliam in the Western Cape

(approximately 1 100 km), and extended inland several hundred kilometres.

Even in this broader context, however, many of Péringuey’s SAM specimens,

including several types, recorded from Namaqualand are likely to bear

erroneous localities and should be viewed with extreme caution (Table 1).

The type of Mabuya peringueyi (= Mabuya homalocephala peringueyi) is

catalogued as collected in Damaraland (although Boulenger (1888) gave the

locality as Namaqualand in the type description). No other specimens of

Mabuya homalocephala are known from north of the Orange River and the

closest modern records are well to the south of Namaqualand, from the Cedar-

berg (Branch 1990a) and the Karoo National Park, Beaufort West (Branch &

Braack 1989).

Similarly, neither of the snakes Duberria lutrix or Crotaphopeltis hotam-

boeia, supposedly obtained in Namaqualand by Péringuey, is found north of

latitude 31°S. Despite its scientific name, Pseudocordylus microlepidotus nama-

quensis Hewitt, 1927, also based on Péringuey material, is not known from

Namaqualand. It is restricted to the Nuweveldberg, which forms part of the

inland escarpment mountains of the Cape (Branch & Braack 1989).

The most striking parallel with respect to Phyllodactylus peringueyi, is the

case of SAM-799, also supposedly from Namaqualand. Although unidentified

when accessioned, it was later identified as Tetradactylus africanus by

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Boulenger (1910) and subsequently as 7. a. fitzsimonsi by FitzSimons (1943).

The latter author regarded the locality as unacceptable and noted (p. 298) that

the ‘specimen in question probably comes from Port Elizabeth’. Indeed, this

subspecies is now known to be restricted to the environs of Port Elizabeth

(Branch 1990b).

TABLE 1

South African Museum herpetological specimens catalogued from Namaqualand during the

same period as Phyllodactylus peringueyi type specimens.

Species SAM Catalogue Known distribution

No. locality

Phyllodactylus peringueyi 777 (syntype) Namaqualand Kromme River estuary

Duberria lutrix 770 Namaqualand Southern Cape, Beaufort West

Crotaphopeltis hotamboeia az Namaqualand Southern Cape, Beaufort West

Agama atricollis 849 Namaqualand Northern Namibia and

Transvaal lowveld

Pseudocordylus microlepidotus 872 (type), 859, | Namaqualand Sutherland to Beaufort West

namaquensis 864, 873

Varanus albigularis 875 Namaqualand __ Eastern southern Africa and

Central Namibia

Pachydactylus fasciatus 1052 (type) Namaqualand Northern Namibia

1155 Natal

Mabuya homalocephala peringueyi 1054 (type) Damaraland No recent specimens

? (later identified as Tetradactylus 799 Namaqualand _— Port Elizabeth

africanus by Boulenger (1910)

and updated to T. a. fitz- _

simonsi by FitzSimons (1943))

Similar locality problems plague other specimens associated with Péringuey.

Another gecko, Pachydactylus fasciatus, was described by Boulenger (1888)

from specimens supposedly collected by Péringuey from Namaqualand. Fitz-

Simons (1943: 88) noted that *. . . there is a specimen (No. 1155) in the South

African Museum, recorded from Natal; this specimen is identical to the cotype,

(in structure, state of preservation etc.) and in all probability came from the

same locality; its occurrence in Natal is otherwise unknown and beyond the

bounds of possibility.” Even the Namaqualand locality ‘between the mouth of

the Orange River and Walfisch (Walvis) Bay’ (Boulenger 1888: 136) is probably

incorrect. The known distribution of Pachydactylus fasciatus is further north,

with the southernmost confirmed record at Neu Barmen, near Windhoek. In

addition, the species appears to be associated with the slopes of the inland

escarpment rather than with coastal desert (Bauer & Branch 1991).

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI V5)

Nothing in the early SAM annual reports indicates that Péringuey ever

visited Namibia, and a number of the specimens attributed to Péringuey in

Boulenger’s 1888 paper were, in reality, collected by Nightingale (e.g. Oedura

(= Afroedura) africana, Pythonodipsas carinata, and Vipera (= Bitis)

peringueyi) or Marloth (Agama planiceps) (Boulenger 1910). Nightingale

donated a considerable amount of material (mainly entomological) to the SAM,

mostly in 1886. It was shipped from Walvis Bay, although much would have

been collected from further inland.

Although no additional specimens of Phyllodactylus peringueyi have ever

been found around Port Elizabeth, there are no historical reasons to doubt the

locality of the paralectotype supposedly collected there by Moorhouse in 1904.

Moorhouse appeared on the Port Elizabeth voters roll for 1905-1907, and is

there listed as a commercial traveller with Blaine and Co. (J. Bennie pers.

comm.). His occupation raises the possibility that he may have collected

specimens elsewhere on his travels, and SAM-8628 may have mistakenly been

accessioned under Moorhouse’s home address. However, this is unlikely as

Chelsea Point, an uninhabited coastal landmark approximately 10 km due south

of Port Elizabeth, is specifically listed.

Moorhouse was not a regular collector for the SAM and there is only one

additional donation from him—a series of 44 specimens, including 3 species of

frog, 8 species of lizard, and 13 species of snake (SAM-9021 to 9026, 9119 to

9146, 9149 to 9159). All the species are currently found in the Port Elizabeth

region (Branch 1988a), adding circumstantial support to the correctness of the

locality for the P. peringueyi paralectotype.

In addition to the problem of locality, there is a problem as to the date of

collection of the syntypes. Hewitt (1937) noted that the nearly identical state of

conservation of the specimens suggested that they had been collected and pre-

served at the same time. This does seem to be the case. Phyllodactylus perin-

gueyi is so obviously different from other southern African geckos (due to its

tubercular dorsal scalation) that it seems unlikely the Namaqualand specimen

would not have been recognized as unusual and taken to Europe by Péringuey

when he visited Boulenger in 1886-1887. It is therefore likely to have been

acquired after 1888, not 1885 as listed. When Boulenger received the Chelsea

Point specimen, after Sclater’s letter in 1904, he immediately recognized its dis-

tinctiveness and described it as a new species (Boulenger 1910). It is therefore

possible that Moorhouse collected both specimens, and that the reference in the

1885 specimen register to Phyllodactylus n. sp. does not refer to SAM-777.

There is no indication that Moorhouse collected more than a single specimen

of Phyllodactylus. None the less, curatorial practice at the SAM at the time gave

only one accession number to all specimens of a species collected at the same

locality. It is most probable that more than one collection was forwarded to the

SAM from Port Elizabeth.

THE DISTRIBUTION OF PHYLLODACTYLUS PERINGUEYI (Fig. 5)

All recent collections of Phyllodactylus peringueyi have been from tran-

sitional salt-marsh vegetation in the lower Kromme River. It has not been found

in neighbouring estuaries or the adjacent coastline. If P. peringueyi was indeed

collected at Chelsea Point, it is possible that it is now extinct there. The Port

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Elizabeth region has been very well surveyed herpetologically, but no new

specimens have been found. The senior author has been based in Port Elizabeth

for over 12 years and has undertaken numerous extensive searches for the spe-

cies. Chelsea Point is located on the south-western boundary of Cape Recife, an

area that consists of shifting coastal dunes with scattered low sandstone outcrops

and regions of calcrete pavement. It is naturally vegetated with a thick, dwarf

heathland (fynbos), but is now heavily infested with introduced alien Australian

Acacia. This was introduced at the turn of the century to control drifting sand

dunes around the environs of the burgeoning city of Port Elizabeth. The dense

stands of exotic Acacia now lead to extensive fires that may have resulted in the

extinction of the species. These plants have been implicated in the reduction in

numbers of the local berg adder, Bitis atropos (Branch 1988b). Alterations in

fire regimes and afforestation with exotic timber plantations are believed to be

responsible for the possible extinction of one other South African lizard, Tetra-

dactylus eastwoodae (Jacobsen 1988).

;

oO Sundays River

ALGOA BAY

Gamt Ri i

Costes Van Staden’s River Swartkops River

PL PORT ELIZABETH

Maitland River

Kabeljous River 2

St Francis Bay \

Chelsea Point

= ONee

if ft Kromme River

INDIAN OCEAN

(0) 30 km

Fig. 5. Distribution of Phyllodactylus peringueyi. Listed collecting localities of the syntypes

are shown by arrows: 1—Little Namaqualand, 2—Chelsea Point. The two arrows on the

Kromme River in the larger map show the current limits of the range of P. peringueyi along

the tidal reaches of the river.

The apparent restriction of P. peringueyi to the Kromme River estuary is

biogeographically problematic. Such low-lying areas rarely harbour endemic

species. Further, the vicissitudes of the estuarine environment suggest that

the species may possess adaptations for coping with hypersalinity and even

sporadic inundation. It is possible that the species has been introduced by human

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 27

agency or more probably by rafting. McLachlan (1988) noted that the Chelsea

Point specimen had been found on the beach and suggested that the animal may

have thus arrived as a result of overwater waif dispersal. Overwater transport

has been implicated in other leaf-toed geckos. For example, the Galapagos

species of Phyllodactylus appear to be derived from three overwater founder

events, each originating from mainland western South America (Wright 1983).

Other American Phyllodactylus are restricted to strand environments (Dixon &

Huey 1970), although such forms have broader distributions than P. peringueyi.

RELATIONSHIPS OF PHYLLODACTYLUS PERINGUEYI

Hewitt (1937) listed a number of differences between Phyllodactylus

peringueyi and other South African Phyllodactylus species, including: the

markedly elongate snout; dark streak on the side of the head that passes through

the eye; the broadly bell-shaped mental; the absence of chin shields or a median

cleft in the rostral. He concluded (p. 206): ‘It seems to agree best with the

American group of species, and I do not accept it as South African.’ This view

was tentatively supported by FitzSimons (1943). Loveridge (1947) reported on

Hewitt’s statements without further comment of his own. The most recent

review of P. peringueyi was provided by McLachlan (1988), who again ques-

tioned the origin of the specimens, although Dixon (in McLachlan 1988) has

ruled out the specific identity of P. peringueyi with either American or

Madagascan leaf-toed geckos, and our results confirm this.

Phyllodactylus peringueyi is a valid species that is morphologically distinct

from all other described leaf-toed geckos. Specific identity with most other leaf-

toed gecko species can easily be ruled out on the basis of external characters,

and many species can be eliminated on the basis of size. The largest P. perin-

gueyi has a SVL of 28.3 mm and radiographs clearly show that all long bone

epiphyses are well ossified. Gravid females have SVLs of 26-28 mm, and the

species is obviously a dwarf form. The greatly enlarged preanal scales typical of

P. gerrhopygus, P. heterurus and P. angustidigitatus (Dixon & Huey 1970;

Kroll & Dixon 1972) and the atuberculate condition of P. leei and P. unctus

(Dixon 1964) are presumably apomorphic conditions within the genus; both

conditions are lacking in P. peringueyi.

Because of the poor condition of the syntypes and the lack of published

comparative data on other Phyllodactylus sensu lato, it has not been possible to

determine the phylogenetic relationships of P. peringueyi. None the less, mem-

bership of the taxon in certain putatively monophyletic groups can be ruled out

on the basis of the lack of one or more apomorphic traits. The nasal bones of

Phyllodactylus peringueyi are paired and there is no evidence of a casque or

other cranial ornamentation. The retention of the plesiomorphic condition for

these features excludes P. peringueyi from membership in Paroedura (Dixon &

Kroll 1974). Likewise, the presence of cloacal bones in males precludes close

affinity with Asaccus, in which cloacal bones are entirely absent. The same

criterion would remove the Socotran species, P. riebeckii and P. trachyrhinus

from consideration as closest relatives (Kluge 1982). Further limitation of the

affinities of the species is hindered by the lack of synapomorphic features

diagnosing subunits within the remaining, polyphyletic assemblage currently

subsumed within Phyllodactylus (Kluge 1983). Karyotypic and morphological

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

data obtained by Gordon (pers. comm.) have suggested a phylogeny for the

South African Phyllodactylus species that tentatively links P. peringueyi to

P. porphyreus, but the analysis included only the South African species. On bio-

geographic grounds it seems likely that P. peringueyi is part of an endemic

southern African radiation of leaf-toed geckos. The monophyly of the group and

the affinities of P. peringueyi within it will be assessed elsewhere in a broader

phylogenetic analysis of African Phyllodactylus (Bauer & Branch, in prep.).

ACKNOWLEDGEMENTS

We are grateful to Mike Cherry (Department of Herpetology, South African

Museum) for forwarding to us the syntypes of Phyllodactylus peringueyi, and

for assistance with the early SAM catalogue and correspondence relating to the

syntypes. Gordon Craig (Port Elizabeth) rediscovered the ‘long-lost’ P. perin-

gueyi, and he and his family helped enthusiastically in searches for the species

in the Kromme River estuary region. Gerald Haagner, Rob Hall, M. Burger and

several other colleagues helped to collect fresh material. Murray Ralfe prepared

the excellent head and foot line-drawings of this minute species. Harold Voris

(FMNH), George Zug (USNM), Jens Vindum (CAS), Colin McCarthy and

Nick Arnold (BMNH), David Good (LSU), John Cadle (ANSP) and William

Duellman (KU) kindly loaned or permitted the examination of specimens in

their care. Ken Tighe (USNM) kindly prepared the X-rays used in this study.

Marcia Field (Villanova University) prepared the X-ray positive prints. Funding

from Villanova University, the California Academy of Sciences, and the John

Ellerman Trust at the University of Stellenbosch supported travel of AMB to

South Africa.

REFERENCES

BAUER, A. M. & BRANCH, W. R. 1991. Notes on Pachydactylus fasciatus, a poorly known

gecko from northern Namibia. Journal of the Herpetological Association of Africa 39:

1-5.

BOULENGER, G. 1888. On new or little-known South-African reptiles. Annals and Maga-

zine of Natural History (6) 2: 136-141.

BOULENGER, G. A. 1910. A revised list of the South African reptiles and batrachians, with

synoptic tables, special reference to the specimens in the South African Museum, and

descriptions of new species. Annals of the South African Museum 5: 455-538.

BRANCH, W. R. 1988a. Field guide to the snakes and other reptiles of southern Africa.

Cape Town: Struik.

BRANCH, W. R. 19885. Terrestrial reptiles and amphibians. Jn: LUBKE, R. A., GEss, F. W.

& BRuTON, M. N. eds. A field guide to the Eastern Cape coast: 251-264. Grahams-

town: The Wildlife Society of Southern Africa.

BRANCH, W. R. (ed.) 1988c. South African red data book—reptiles and amphibians. South

African National Scientific Programmes Report 151: i-vi, 1-241.

BRANCH, W. R. 1990a. The herpetofauna of the Cape Province, South Africa: new distribu-

tion records and zoogeography. Journal of the Herpetological Association of Africa 37:

17-44.

BRANCH, W. R. 199056. The genus Terradactylus (Sauria: Gerrhosaurinae) in the Cape Pro-

vince, South Africa: new records and their taxonomic status. Journal of the Herpetol-

ogical Association of Africa 37: 13-16.

REDISCOVERY OF PHYLLODACTYLUS PERINGUEYI 29

BRANCH, W. R. & BAueR, A. M. 1992. The status of Phyllodactylus peringueyi (Reptilia:

Gekkonidae). (Extended abstract). Jn: BRANCH, W. R., HAAGNER, G. V. & BoycoTrT,

R. C. eds. Proceedings of the Second Herpetological Association of Africa Symposium.

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BRANCH, W. R. & Braack, H. H. 1989. Reptiles and amphibians of the Karoo National

Park: a surprising diversity. Jn: BRANCH, W. R. ed. Proceedings of the First Herpetol-

ogical Association of Africa Conference, Stellenbosch. Journal of the Herpetological

Association of Africa 36: 26-35.

BRANCH, W. R., HAAGNER, G., HALL, R. & CraiGc, G. 1992. Phyllodactylus peringueyi:

geographical distribution. Journal of the Herpetological Association of Africa 41: 38.

CoccER, H. G. 1986. Reptiles and amphibians of Australia. 4th ed. Frenchs Forest: Reed

Books Pty. Ltd.

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in North and Central America. Bulletin. Research Center, New Mexico State University

64-1: i-iv, 1-139.

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McLacHLAaN, G. 1988. Péringuey’s leaftoed gecko Phyllodactylus peringueyi. In: BRANCH,

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organisms: 123-155. San Francisco: Pacific Division, American Association for the

Advancement of Science.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

APPENDIX

Material of Phyllodactylus peringueyi examined:

SAM-777, Little Namaqualand, Cape Province, South Africa (lectotype).

SAM-8628, Chelsea Point, near Port Elizabeth, Cape Province, South Africa

(paralectotype).

PEM R6886, 6916, 7210, Kaia da Balaia, Kromme River, Humansdorp, Cape

Province South Africa (3424Bb; 34°07'15"S 24°45'30"E; alt. 1 m asl.).

PEM R6908-6915, region of road bridge at mouth of Kromme River (3424Bb;

34°08'15"S 24°48'50"E).

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particu-

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. ‘.. . the Figure depicting C. namacolus ...’: ‘. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by

initials or full names

e.g. DuToit but A.L. du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

not be abbreviated at the beginning of a sentence or paragraph.

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.