European Journal of Taxonomy 194: 1-16

http://dx.doi.org/10.5852/ejt.2016.194

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This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Roman-Palacios C. et al.

Research article

urn:lsid:zoobank.org:pub:96E9EA43-F6FE-492E-97BE-60DFB8EDE935

Historical biogeography of Thyrsophorini psocids

and description of a new neotropical species of Thyrsopsocopsis

(Psocodea: Psocomorpha: Psocidae)

Cristian ROMAN-PALACIOS '•*, Alfonso N. GARCIA ALDRETE 2 &

Ranulfo GONZALEZ OB ANDO 3

13 Departamento de Biologia, Facultad de Ciencias Naturales y Exactas, Universidad del Valle,

Santiago de Cali, Colombia.

2 Departamento de Zoologia, Instituto de Biologia, Universidad Nacional Autonoma de Mexico,

Apartado Postal 70-153, 04510 Mexico City, Mexico.

* Corresponding author: [email protected]

1 um:lsid:zoobank.org:author:E88D0518-B6CB-4FE7-9EFC-F789EA6F05AD

2 um:lsid:zoobank.org:author:9E03B921-78AE-4ED6-BlEA-9DCA01BE20BC

3 urn:lsid:zoobank.org:author:16C7AD76-F035-4C8B-8C00-A228CCCD39B0

Abstract. When based on phylogenetic proposals, biogeographic historic narratives have a great

interest for hypothesizing paths of origin of the current biodiversity. Among the many questions that

remain unsolved about psocids, the distribution of Thyrsophorini represents still a remarkable enigma.

This tribe had been considered as exclusively Neotropical, until the description of Thyrsopsocopsis

thorntoni Mockford, 2004, from Vietnam. Three hypotheses have been proposed to explain this atypical

distribution, recurring to dispersal, vicariance and morphological parallelism between lineages, but

the lack of evidence has not allowed a unique support. Here, we describe a new Neotropical species

of Thyrsopsocopsis , and also attempt to test the three biogeographical hypotheses in a phylogenetic

context. In our analyses, the position of Cycetes (Cycetini) and Thyrsopsocopsis among Thyrsophorini

psocids is well resolved, suggesting a dispersal event from the Neotropics for both genera.

Keywords. Historical biogeography, Psocids, dispersal, vicariance, parallelism.

Roman-Palacios C., Garcia Aldrete A.N. & Gonzalez Obando R. 2016. Historical biogeography of Thyrsophorini

psocids and description of a new neotropical species of Thyrsopsocopsis (Psocodea: Psocomorpha: Psocidae).

European Journal of Taxonomy 194: 1-16. http://dx.doi.org/10.5852/ejt.2016.194

Introduction

Psocidae is the largest and most diversified family of psocodeans (Lienhard & Smithers 2002). Most

of what has been published on the family consists of taxonomic papers, and little has been published

on evolutionary or biogeographic aspects of it. The inclusion of molecular data on recent phylogenetic

analyses has helped much to clarify the position of some genera and their status (Yoshizawa &

Johnson 2008, Yoshizawa et al. 2011), but its limited availability predisposes the family to a major

morphologically oriented work (Li 2002).

1

European Journal of Taxonomy 194 : 1-16 ( 2016 )

The classification of the various suprageneric lineages within Psocidae has varied through time, mainly

due to the high morphological complexity (Lienhard 2008; Yoshizawa & Johnson 2008). Currently,

three subfamilies are recognized: Kaindipsocinae, Amphigerontiinae and Psocinae (Yoshizawa et al.

2011), the latter including five tribes among which Sigmatoneurini, Cycetini and Thyrsophorini are

remarkable by their peculiar morphological plan (New 1978; Mockford 1992). Cycetini is characterized

by a particular forewing venation along with the structure of female gonapophyses (Roesler 1940;

Smithers 1972), but its position and systematic validity has not been tested within any phylogenetic

scheme. The ambiguous status of this tribe within Psocinae has made impossible to establish a clear

phylogenetic reference for discussing biogeographical or evolutionary aspects of other lineages as

Thyrsophor ini (Mockford 2004).

Thyrsophorini includes two major groups, previously referred as Cerastipsocinae and Thyrsophorinae

(Yoshizawa & Johnson 2008). This tribe, that includes the largest psocids (Mockford 2004, Smithers

1972), groups 15 widely distributed genera. Thyrsopsocus Enderlein, 1900, Thyrsophorus Burmeister,

1839, Dictyopsocus Enderlein, 1901, Poecilopsocus Roesler, 1940, Eremopsocus McLachlan, 1866,

Cervopsocus New, 1978 and Ghesquierella Badonnel, 1949 are exclusively Neotropical lineages.

Thyrsopsocopsis Mockford, 2004, along with Setopsocus Smithers & Thornton, 1981, have been collected

in the Oriental region; Cerastipsocus Kolbe, 1884 is the only Neartic-Neotropical genus. Neopsocus

Kolbe, 1882, Longivalvus Li, 1993 and Sundapsocus Smithers, 1995 are distributed in the Palearctic.

Finally, the specious Psococerastis Pearman, 1932 is widely distributed and Clematoscenea Enderlein,

1907 is restricted to the Oriental, Palearctic and Australasian regions (Lienhard & Smithers 2002).

The genus Thyrsopsocopsis was erected by Mockford (2004) based on one male specimen collected

in Vietnam and assigned to the subfamily Thyrsophorinae (currently Thyrsophorini), mainly by its

morphological resemblance to the Neotropical Thyrsophorus. As Mockford (2004) pointed out,

the establishment of the new genus within this exclusive Neotropical lineage represents “a difficult

biogeographical problem”, which can only be elucidated from an evolutionary context and by taking

into account multiple aspects of the natural history of the tribe.

Several plausible explanations have been proposed to deal with the atypical distribution of this genus

in relation to other Thyrsophorini. Mockford (2004) argues that this distribution could be consequence

of (i) a modern dispersal to the Oriental region, as has been indicated for other psocids (Fig. 1A). This

hypothesis implies a large specific richness of Thyrsopsocopsis in the Neotropics, and the sympatry of

several (currently unknown) species along with T. thorntoni Mockford, 2004 in the same region. An

alternative hypothesis suggests an (ii) ancient vicariance generated by the separation of the tropical

lands (Fig. IB). This event, which might have occurred near the Permian, establishes the possibility

of the splitting of populations and subsequent speciation of the most recent common ancestor between

Cycetini and Thyrsophor ini This approach implies Cycetini more diverse than Thyrsophor ini in earlier

times, following the centers of dispersal proposal. Finally, the possibility of (iii) parallel evolution

between Neotropical and Oriental lineages, as a consequence of similar environmental conditions

(Fig. 1C) has been also considered. In this paper we describe a new species of Thyrsopsocopsis from

Brazil, propose a generic-level morphological phylogeny for Thyrsophorini, and discuss the systematics

and biogeographical-evolutionary aspects of the tribe.

Material and methods

Taxonomy

The two Brazilian individuals available of Thyrsopsocopsis were dissected in 80% ethanol. The

head, terminalia, right wings and legs were mounted on slides in Canada balsam. Measurements and

photographs were taken using an AxioCam MRc5 mounted on an Axio zoom vl6 microscope and

2

ROMAN-PALACIOS C. et al. Historical biogeography of Thyrsophorini

stacked using the ZEN software (Zeiss Efficient Navigation). Color was recorded by placing the entire

specimens under a microscope illuminated with cold white light at 40X. The illustrations were processed

in the vector graphics editor CorelDraw X7.

B

11

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Remaining

Psocidae

Remaining

Psocinae

Thyrsophorini

Fig. 1 . Hypotheses for the current distribution of Thyrsophorini. A. Dispersal of Thyrsopsocopsis

Mockford, 2004 from the Neotropics. B. Vicariance, as a consequence of the population split, occurred

in the hypothesized most recent co mm on ancestor of Thyrsophorini and Cycetini. C. Parallel evolution

between the Neotropical and Asian lineages.

3

European Journal of Taxonomy 194: 1-16 (2016)

Abbreviations for lengths of parts measured are as follows:

bp = basal prominence of mdp (see Mockford, 2004)

D, d = antero-posterior and transverse diameters, respectively, of right compound eye, in dorsal

view of head

fl... fn = flagellomeres 1... n of right ante nn a

FW = right forewing

HW = right hindwing

10 = minimum distance between compound eyes

ldp = latero-distal processes of the hypandrium

mdp = medio-distal processes of the hypandrium

Mx4 = fourth segment of right maxillary palpus

pp = paired proximal processes of the hypandrium

The types will be deposited in the Colesao de Invertebrados, Instituto Nacional de Pesquisas da

Amazonia, Manaus, Amazonas, Brazil.

Hypothesis testing

Systematics

The matrix was constructed in MESQUITE 3.04 (Maddison & Maddison 2008), resulting in 43 non-additive

characters from 27 taxa (Appendices 1-2). The morphological data was analyzedunderparsimony implemented

in TNT (Tree Analysis Using New Technology, Goloboff et al. 2003) using a heuristic search (1000 replicates)

of random addition sequence followed by TBR branch swapping. Characters were weighted using implied

weighting with a concavity constant (k) from 1 to 6, selecting a k = 3. Node support was estimated using (i)

relative non-parametric bootstrapping, (ii) symmetric resampling using 1000 replicates, and a heuristic search

with a change probability of 33%; these are reported as frequency differences between groups present and

contradicted (GC), and (iii) Bremer support (BS; Bremer 1988, 1994). We selected multiple outgroups for

testing the position of Cycetini and the monophyly of Thyrsophorini in accordance to Yoshizawa & Johnson

(2008): Amphigerontia Kolbe, 1880 (Amphigerontiinae), Elaphopsocus Roesler, 1940 (Amphigerontiinae),

Elaphopsocoides Roman-P, Garcia Aldrete & Gonzalez-Obando, 2014 (Amphigerontiinae), Blastopsocus

Roesler, 1943 (Amphigerontiinae), Ptycta Enderlein, 1925 (Psocinae), Trichadenotecnum Enderlein, 1909

(Psocinae), Atrichadenotecnum Yoshizawa, 1998 (Psocinae), Metylophorus Peaiman, 1932 (Psocinae),

Elyalopsocus Roesler, 1954 (Psocinae) and Sigmatoneura Enderlein, 1908 (Psocinae).

Historical biogeography

To include the biogeographic context in the phylogenetic interpretation of the Thyrsophorini, we used

the ancestral reconstruction implemented in MESQUITE 3.04 (Maddison & Maddison 2008) and

RASP software (Yu et al. 2015). The former uses unweighted parsimony for tracing the character states

through the tree and RASP evaluates the hypothesized ancestral ranges at each node in the tree by

using a statistical framework, thus accounting for uncertainties both in phylogenetic inference and in

biogeographic optimization (Pialek et al. 2012). The S-DIVA (Statistical Dispersal-Vicariance Analysis,

modified from DIVA) and BBM (Bayesian Binary MCMC) in the package RASP were used to reconstruct

the ancestral geographic distributions. The areas used and the distributions of the sampled species are

given in Fig. 5 A. The analysis uses distributions of contemporary species, which does not mean assuming

similar continental configurations through time. The MCMC (Markov chains Monte Carlo) analyses in

BBM were based on 50000 states, with a 10% burn-in generation. All biogeographical reconstructions

were based in Maximum Likelihood (ML) topology, obtained from RAxMLGUI analyses (Silvestro

& Michalak 2011) by means of a MULTIGAMMA + Mk model, implementing 1000 replicates for the

bootstrap support and selecting a single outgroup (< Sigmatoneura ), to limit the uncertainty that widely

distributed and poorly known taxa would imply for the analyses; other taxa were pruned using the

package APE (Paradis et al. 2004) in R software (R Core Team 2013).

4

A

ROMAN-PALACIOS C. et al ., Historical biogeography of Thyrsophorini

Morphological parallelism

The hypandrium was evaluated in order to account for the more variable structure in the two species

of Thyrsopsocopsis , and test the morphological parallelism between Asian and Neotropical lineages.

Therefore, it’s assumed that morphological resemblance would suggest a narrow phylogenetic relation

between taxa. The anatomical homology of the hypandrial morphological structure was assessed by

external examination. Other characters commonly used for morphological phylogenetic analyses

(e.g., phallosome or forewing morphology) are remarkably similar between the two species (compare

Fig. 2A-F in this document with figures presented in Mockford, 2004).

Results

Taxonomy

Order Psocodea Hennig, 1966

Suborder Psocomorpha Badonnel, 1951

Family Psocidae Hagen, 1865

Genus Thyrsopsocopsis Mockford, 2004

Thyrsopsocopsis amazonicus sp. nov.

um:lsid:zoobank.org:act:87C7ElC7-164C-461E-95D2-38F5B77B907D

Fig. 2A-F

Diagnosis

As in generic diagnosis (see Mockford 2004), plus the following: clunium bearing on the posterior

margin a pair or heavily sclerotized lateral spines. Paraprocts with proximal handle. Hypandrium with

pp, mbp and ldp anteriorly directed (posteriorly directed in T. thorntoni); bp absent.

Etymology

The specific name refers to the nature of the area ( Amazonian forest), where the types were collected.

Type material

Holotype

BRAZIL: c?, Amazonas, AM 010, Reserva Florestal Adolpho Ducke, Mixed light, 2 May 2002, J. Vidal.

Paratype

BRAZIL: 1 same data as the holotype (INPA).

Description (male)

Color (in 70% ethanol). Body light brown. Compound eyes black, ocelli hyaline. A broad dark brown

band over most of the frontal suture, ending at the antennal fossae. Flagellum with fl black, other

flagellomeres brown. Pronotum and mesonotum with median black stripe. Legs brown. Fore- and

hindwings with variable pigmentation and dark brown veins except for Cu la and IA (white); pterostigma

brown; a dark brown spot from discoidal cell to distal area. Hindwing without markings except basally.

Morphology. Compound eyes relatively small, the anterior borders not reaching the level of the convex

vertex; antennae with first flagellomeres thickened and densely covered with setae. Forewing with

elongate pterostigma; clunium bearing a pair of well-developed and heavily sclerotized lateral spines and

with clunial shelf covering base of epiproct. Epiproct simple, rounded and semi-membranous. Posterior

projection of paraprocts long and acuminate, inwardly directed; large sensory fields with many (> 100)

trichobothria in basal rosettes. Hypandrium symmetrical, bearing short and acute median pp; mdp a

5

European Journal of Taxonomy 194: 1-16 (2016)

massive bulb lacking bp; pp distally quadrangular and outwardly directed, bearing spines, spicules and

setae over the surface; a heavily sclerotized and continuous band between pp and mdp. Phallosome closed,

elongate, with a denticulate apical process, ending in a short tooth; endophallus laterally spinulose.

Measurements (in pm). FW: 6600, HW: 4600, Mx4: 330, fl: 740, 12: 2320, f3: 109, IO: 970, D: 210,

d: 350, IO/d: 4.61, PO: 1.6.

Fig. 2. Thyrsopsocopsis amazonicus sp. nov., holotype. A. Front view of head. B. Paraprocts, epiproct

and clunium. C. Forewing. D. Hindwing. E. Phallosome. F. Hypandrium. Scales in mm

6

ROMAN-PALACIOS C. et al ., Historical biogeography of Thyrsophorini

Hypothesis testing

Systematics

The relations among suprageneric groups were reconstructed and highly supported (Fig. 3). Sigmatoneurini

represent the least derived tribe among thyrsophorinae-like lineages. Thyrsophorini was recovered as a

monophyletic lineage, grouping genera previously assigned into separated two lineages (Cerastipsocinae

' Amphigerontia

14

7 11 132026

Elaphopsocus

Elaphopsocoides

Blastopsocus

3740

Ptycta

26

Trichadenoctenum

—o#-

Atrichadenotecnum

i i

Hyalopsocus

14

62 / 31 /

4 114142

1 —o-o#-#

' Metylophorus

172124262840

Pearmania

2126 30

r <K># Sigmatoneura

Eremopsocus

Ghesquierella

11 22242526

Cycetes

Thyrsopsocus

Thyrsophorus

Dictyopsocus

4 12242535

Poecilopsocus

Thyrsopsocopsis

Psococerastis

8 23

4 34 Neopsocus

Sundapsocus

Clematoscenea

Longivalvus

Fig. 3. Generic-level cladogram derived from parsimony analysis of the morphological dataset based on

a heuristic search with 1000 replicates (L = 127, Cl = 40, Ri = 59), followed by TBR branch swapping,

with character optimizations, symmetric resampling (*0.33), Bremmer support (*greater than 0.33 for

the current dataset) and relative bootstrap (*greater than 70%). Circles represent character optimizations:

o = unique changes; • = homoplasic characters.

7

European Journal of Taxonomy 194: 1-16 (2016)

and Thyrsophorinae). The morphological phylogeny also indicated Cycetini ( Cycetes Enderlein, 1907)

merged within Thyrsophorini, suggesting a narrow relation between both lineages. The phylogenetic

hypothesis supports two of the three hypotheses, but cannot differentiate between them: (i) a modern

dispersal even for the Neotropical region and (ii) a vicariance event that involved the most recent common

ancestor of both lineages. The parallelism proposal is not supported by the phylogenetic relationships

recovered, because of the position of Cycetes within Thyrsophorini.

Historical biogeography

Historical reconstructions generated by parsimony, statistical parsimony and Bayesian algorithms,

indicated similar and comparable tendencies in the biogeography of Thyrsophorini. The reconstruction of

ancestral characters by unweighted parsimony indicated an ancestral Neotropical distribution for the tribe,

and, consequently, the same distribution for the Cycetes ancestor (Fig. 4A). The genus Thyrsopsocopsis

appeared as a lineage derived from a Neotropical ancestor. The current biogeographical evidence allows

inferring the monophyly of a Palaearctic lineage, corresponding to the most recent co mm on ancestor

(MRCA), between Sundapsocus and Clematoscenea. The genus Psococerastis corresponds to a lineage

with wider geographical distribution, but the analysis indicates the equivalent probability of an ancestral

Neotropical or Palaearctic distribution. Statistical parsimony showed uncertainty in the ancestral

distribution of most nodes in Thyrsophorini phylogeny (Fig. 4B). The MRCA between Thyrsopsocopsis

and Poecilopsocus showed an exclusively Neotropical distribution.

Bayesian reconstruction of the historical biogeographic distribution of Thyrsophorini presented a major

resolution in the internal nodes (Fig. 5). The ancestral distribution for Thyrsophorini resulted exclusively

Neotropical, which agrees with the results of previous analyses. Thyrsopsocopsis represents a modern

dispersal towards the Oriental region (T. thorntoni). The consensus analysis of the reconstructions

obtained in S-DIVA and BBM (Fig. 6) showed a notable similarity in the results, and hence congruence

between independent reconstructions. Cycetes may correspond to a dispersal from the Neotropics

towards the Oriental-Australasian region, with a subsequent vicariance, resulting in the two species

known for each region (C. collesi Smithers, 1977, C. thyrsophoroides Enderlein, 1907). An initial

vicariance probably in the MRCA of Clematoscenea and Setopsocus generated the divergence between

Neotropical and Palaearctic ancestors; the MRCA in Sundapsocus and Setopsocus corresponded to

a dispersal from the Oriental region. Sundapsocus and Cervopsocus possibly diverged by vicariance

among Neotropical and Palaearctic ancestral populations, similar to what occurred in the MRCA of

Setopsocus and Clematoscenea. The principal biogeographic trend in Thyrsophorini has been one of

dispersal from the Neotropics to the Oriental region (4 events), with the first region having the largest

number of variance events within the area (9 events).

Morphological parallelism

This hypothesis proposes parallel morphologies in both lineages (Fig. 7), which can be contrasted

from the more variable structure between the two species. The hypandrium of T. thorntoni bears a

basal projection (bp), lacking in T. amazonicus sp. nov., but the background morphological plan of the

hypandrium of both species is well conserved. Clunial projections found on males of T. amazonicus

sp. nov. are absent in T. thorntoni. Finally, the phylogenetic position of Cycetini within Thyrsophorini

does not support this hypothesis (see above).

Discussion

The former Thyrsophorini had been considered as a lineage restricted to the Neotropics, with a possible

origin from Cycetini-like ancestors in the Oriental region (Mockford 2004). Since the inclusion of

Cerastipsocini within Thyrsophorini (Yoshizawa & Johnson 2008), the biogeographic heterogeneity

has ceased to be a valid reference for the identification of these taxa, moreover, the description of

8

ROMAN-PALACIOS C. et al ., Historical biogeography of Thyrsophorini

LEGEND

*

Fig. 4. Parsimony based reconstruction of Thyrsophorini historical biogeography. A. Parsimony based

ancestral reconstruction implemented in Mesquite 3.04. B. S-DIVA biogeographical reconstruction.

9

European Journal of Taxonomy 194: 1-16 (2016)

LEGEND

*

Sigmatoneura

Ghesquierella

Eremopsocus

Cerastipsocus

Clematoscenea

Longivalvus

Neopsocus

Psococerastis

Sundapsocus

Cervopsocus

Setopsocus

Cycetes

Thyrsopsocus

Poecilopsocus

Thyrsopsocopsis

Dictyopsocus

Thyrsophorus

Fig. 5. Bayesian algorithms based reconstruction of the Thyrsophorini biogeography. A. Biogeographic

regions considered in the analyses: Blue: Neotropics (A), purple: Oriental (B), Cream: Nearctic (C),

Orange: Paleartic (D), Green: Australasia (E). B. Reconstructed distribution using Bayesian Binary

MCMC (BBM).

10

A

ROMAN-PALACIOS C. et al. , Historical biogeography of Thyrsophorini

Sigmatoneura

Ghesquierella

Eremopsocus

Cerastipsocus

Clematoscenea

Lortgivalvus

Neopsocus

Psococerastis

Sundapsocus

Cervopsocus

Setopsocus

Cycetes

Thyrsopsocus

Poecilopsocus

Thyrsopsocopsis

Dictyopsocus

Thyrsophorus

Fig. 6. Major biogeographical events occurred in Thyrsophorini (left) in accordance to the consensus

reconstruction between BBM and S-DIVA analyses (right). Significant reconstructions indicated with *.

I. Vicariance of the most recent common ancestor between Longivalvus Li, 1993 and Clematoscenea

Enderlein, 1907. II. Allopatric speciation in the MRCAof Sundapsocus Smithers, 1995 and Cervopsocus

New, 1978. III. Dispersal event occurred in Thyrsopsocopsis Mockford, 2004.

P

A

Fig. 7. Morphological homology of the hypandrial tubercles between the Neotropical and Oriental

species. A. T. amazonicus sp. nov. B. T. thorntoni Mockford, 2004. Anterior -A and -P axis indicated

on left. Scales in mm.

11

European Journal of Taxonomy 194: 1-16 (2016)

Thyrsopsocopsis shed doubts about the current knowledge on the evolutionary and biogeographic

aspects of Thyrsophorini (Yoshizawa & Johnson 2008). Thyrsopsocopsis thorntoni represented a change

in the paradigm of the distribution of New World Thyrsophorini, allowing for hypothesizing different

scenarios in the evolution of this clade (see Moclcford 2004).

Cycetini has been given a tribal status for its remarkable morphological and biogeographic peculiarities

with respect to the described thyrsophorines, but the description of the two known species of Cycetes was

based on females, which do not provide an objective evidence for a correct assignation within Psocidae

(New & Lienhard 2007). The taxonomy of the family has been based mostly on male characters, and the

male-female associations have been made on ecological and/or temporal evidences (e.g., Kaindipsocinae;

Yoshizawa et al. 2011). Despite this, phylogenetic analyses carried out in the present work indicated that,

although the wing and genital morphology of Cycetes are different from that usually found in Thyrsphorini,

they represent generic apomorphies, brushing aside their consideration as tribal synapomorphies.

The systematic position of Cycetini in Thyrsophorini questions its tribal status. The phylogeny of

Thyrsophorini agrees with the phylogenetic proposal made by Yoshizawa & Johnson (2008), supporting the

monophyly of Thyrsophorini, but resulted in several remarkable differences (e.g., compare Thyrsopsocus +

Cerastipsocus in Yoshizawa & Johnson 2008), which could be related to (i) the high morphological homoplasy

found in Thyrsophorini morphology or (ii) a consequence of gene-tree based methods used in molecular

analyses. The phylogenetic explanation made by Mockford (2004) is not congruent with that proposed in

the current reconstruction. The author found support for the reciprocal monophyly of Cerastipsocinae and

Thyrsophorinae, which was a clue for recognizing an Asiatic ancestry for the Neotropical thyrsophorines.

The present biogeographic evidence and the position of Cycetini within Thyrsophorini support a Neotropical

origin of this clade. Then, Cycetini represents an event of dispersal with a subsequent vicariance that resulted

in the current two current species, which inhabit Australasian and Indomalayan regions.

Biogeographical reconstruction of the most recent common ancestor between Thyrsopsocopsis and

Poecilopsocus suggest a possible dispersal event in Thyrsopsocopsis. This scenario increases its likelihood

when considering biological (e.g., storms, drifting) or anthropic factors (e.g., commerce, transports) that

could lead to the current distribution. Although the small degree of vagility of psocids is recognized by then-

anatomical fragility (Thornton 1964; Thornton & Harrell 1965), their association with plants could be a

key factor for dispersal, even at a transcontinental level. The description of Thyrsopsocopsis amazonicus

sp. nov., and the richness of Thyrsophorini in the Neotropics makes the presence of Thyrsopsocopsis

thorntoni in the same region quite plausible, but also suggests a higher Neotropical richness for the genus.

The morphology of Thyrsopsocopsis amazonicus sp. nov. shows a remarkable similarity with the

Vietnamese species T. thorntoni. The pattern of wing venation, color, genital structure, and projections in

paraprocts and hypandrium, allow for the assignment of the two species to the same genus (Mockford 2004).

The morphological homology for the Neotropical and Asiatic lineages is also verified from anatomical

evidence. The possibility of parallelism of the Thyrsopsocopsis lineages is falsified by (i) morphological

homology and (ii) phylogenetic monophyly. Therefore, we conclude that the distribution of T. thorntoni

corresponds to an event of modern dispersal, with a potential specific richness higher for Thyrsopsocopsis

in the Neotropics. The tribal status of Cycetini is doubtful by its position within Thyrsophorini, but a

nomenclatural act requires high experimental strictness for the objective establishment of the new clade.

Acknowledments

CRP and RGO are grateful to the Departamento de Biologia, Facultad de Ciencias Naturales y Exactas,

Vicerectoria de Investigaciones, Universidad del Valle, and GIE for research support. ANGA thanks

Instituto de Biologia, Universidad Nacional Autonoma de Mexico, for continuous research support. The

authors thank Dr. K. Yoshizawa and an anonymous reviewer for corrections and interesting discussions.

12

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ROMAN-PALACIOS C. et al. , Historical biogeography of Thyrsophorini

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Manuscript received: 18 October 2015

Manuscript accepted: 13 January 2016

Published on: 6 May 2016

Topic editor: Koen Martens

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark.

14

A

ROMAN-PALACIOS C. et al., Historical biogeography of Thyrsophorini

Appendix 1. Characters and character states.

1. First flagellomere incrassate: (0) Absent; (1) Present.

2. Second flagellomere incrassate: (0) Absent; (1) Present.

3. Veins R4+5 and M: (0) Fused; (1) Free; (2) Joined by a crossvein.

4. Phallosome distally: (0) Open; (1) Closed.

5. Distally closed phallosome morphology: (0) Oval; (1) Subtriangular.

6. Parameres basally: (0) Fused; (1) Free.

7. Chair-shaped epiproct: (0) Absent; (1) Present.

8. Distally ornamented phallosome: (0) Absent; (1) Present.

9. Glandular setae, especially on head: (0) Absent; (1) Present.

10. Cell M3 morphology: (0) Narrow; (1) Normal.

11. Pterostigmal morphology: (0) Elongate; (1) L = A; (2) 2L > A.

12. Areola postica: (0)A>L; (1)A = L.

13. Ramified veins in the central portion of forewing: (0) Absent; (1) Present.

14. Crossvein between areola postica and M: (0) Absent; (1) Present.

15. Hypandrium bearing processes: (0) Absent; (1) Present.

16. Number of processes on hypandrium: (0) One; (1) Two; (2) Three; (3) Four; (4) Six.

17. Distal denticles on hypandrium: (0) Absent; (1) Present.

18. Posterior margin of clunium: (0) Simple; (1) Ornamented.

19. Fore tibia expanded: (0) Absent; (1) Present.

20. Forewing bearing setae: (0) Absent; (1) Present.

21. R2+3 and R4+5 diverging at an angle of: (0) 60°; (1) 90°.

22. Distal margin of forewing: (0) Acute; (1) Rounded.

23. Ventral prolongation of VI: (0) Absent; (1) Present.

24. Short setae over v2: (0) Absent; (1) Present.

25. Forewing with Sc ending: (0) Free; (1) in costal vein; (2) in R vein.

26. Hindwing with Sc ending: (0) Free; (1) in costal vein; (2) in R vein.

27. Subgenital plate pigmentation: (0) T-shaped; (1) V-shaped; (2) Different.

28. Hypandrium: (0) Symmetrical; (1) Asymmetrical.

29. Gonapophyses with v3: (0) Normal; (1) Well developed.

30. R4+5 sigmoidal: (0) Absent; (1) Present.

31. Extension of the clunial shelf over the epiproct: (0) Absent; (1) Present.

32. Discal cell in longitudinal position: (0) Absent; (1) Present.

33. Male paraproct bearing a distal projection: (0) Absent; (1) Present.

34. Maxillary palpomere IV: (0) Large; (1) Short.

35. Rs-M: (0) Joined by a crossvein; (1) Fused.

36. VeinM+Cula: (0) Straight; (1) Sinuous.

37. Basal lobe on male paraproct: (0) Absent; (1) Present.

38. Opposite points over cell r: (0) Absent; (1) Present.

39. Triangular areola postica: (0) Present; (1) Absent.

40. Median strap on hypandrium: (0) Absent; (1) Present.

41. Gonapophyses with V2: (0) Slim; (1) Robust.

42. Nimphal aggregation: (0) Absent; (1) Present.

43. Forewing length: (0) Less than 4 mm ; (1) More than 4 mm

15

Appendix 2. The first state is “0”, followed by “1”, etc.; “?” = missing data; = not applies.

European Journal of Taxonomy 194: 1-16 (2016)

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Thyrsophorus

Dictyopsocus

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Cerastipsocus

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Clematoscenea

Ghesquierella

Neopsocus

Cervopsocus

Sundapsocus

Longivalvus

Cycetes

Metylophoms

Sigmatoneura

Pearmania

Amphigerontia

Elaphopsocus

Elaph opsocoides

Blastopsocus

Ptycta

Trick aden otecnum

Hyalopsocus

Atricliadenotecnum

16