Cimbabesia. 2003

Distributional patterns of neotropical, afrotropical and australian-oriental

Simulium Latreille subgenera (Diptera: Simuliidae)

Daniel Rafael Miranda-Esquivel1 & Sixto Coscarón2 1Laboratorio de Sistemática y Biogeografía, Escuela de Biología, Universidad

Industrial de Santander, A. A. 678, Bucaramanga. Colombia; e-mail:

dmiranda@uis.edu.co 2Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata,

Paseo del Bosque 1900, La Plata, Argentina; e-mail:

coscaron@museo.unlp.edu.ar

The historical biogeography of Simulium Latreille subgenera of the Neotropical, Afrotropical, and Australian-oriental Regions is reconstructed applying dispersal-vicariance analysis. The results suggest that the genus originated within an area represented by Pangea. This places the origin for the genus in the middle of the Jurassic Period (180-160 Mya).

INTRODUCTION

Simulium Latreille, 1802, is the most speciose simuliid genus, comprising

almost 75% of the described species of the family Simuliidae (Crosskey &

Howard 1997). The genus occurs in all biogeographical regions, although each

subgenus is confined to one or more regions. The first uncontroversial fossil

assigned to the family is a member of the tribe Prosimuliini (sensu Currie 1988),

from the Jurassic/Cretaceous transition, namely: Kovalevimyia lacrimosa

Kalugina, 1991, from Siberia (Currie & Grimaldi 2000). The oldest Southern

Hemisphere fossil originates from the Cretaceous Koonwarra fossil beds of

Australia (135 Mya), and Crosskey & Howard (1997), assigned this species to

the African genus Paracnephia Rubtsov, 1962, indicating a Gondwanian origin.

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Craig et al. (2001) presented a biogeographic analysis of the subgenus S.

(Inseliellum) Rubtsov, 1974, and suggested that Simulium ‘evolved’ prior to the

separation of New Zealand from Australia/Antarctica (between 80-60 Mya).

Crosskey (1990: 58), believed that the family was widespread in Pangea, and

dated the origin of the family to at least the early Jurassic (200-175 Mya). He

considered, however, dispersal to be the main mechanism explaining the

current distribution patterns of simuliid genera and subgenera without vicariance

playing a crucial rôle. He also proposed that Antarctica would have been

habitable by simuliids at that time, and therefore the continent could have acted

as a bridge between Australia and New Zealand, and South America.

Coscarón & Coscarón-Arias (1995) presented a preliminary biogeographical

analysis of the neotropical simuliid fauna, delimiting 16 areas of endemism.

These provinces, in a slightly modified form, were later used in a

biogeographical analysis by Miranda-Esquivel (2001). In that study the rôle of

dispersal in the distribution of neotropical simuliid subgenera was evaluated.

Eliminating the commonest dispersal events, the general area cladogram was

found to be congruent with terrestrial biota, where two sub-regions arise: the

Neotropical Subregion sensu stricto (Cerrado, SE Brasil, Guyana and

Amazonia, which is synonymous with the Guyano-Brasilian Region of Jeannel

(1942)), and the Andean Sub-region, which is equivalent to the Andean Region

proposed by Morrone (1996). Pacifica and the Mesoamerican Mountains were

placed apart from the main Neotropical Region clade (vide Figure 1). Miranda-

Esquivel & Coscarón (submitted) present a preliminary attempt at a cladistic

analysis of the genus at the subgeneric level for the

Neotropical/Afrotropical/Australian-Oriental Regions.

Event-based methods have increased in importance in historical biogeographic

studies over recent years. Unlike most other pattern-based methods, event-

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based methods postulate explicit models of processes that effect the

geographic distribution of living organisms. In this context, different process

types (i.e. vicariance, dispersal, extinction) are identified, and costs are

assigned in accordance to the model (Ronquist 1996, 1997, 2002). Dispersal-

vicariance analysis (Ronquist 1997), allows for the reconstruction of ancestral

distributions, maximises vicariance events, and minimises dispersal and

extinction events, but unlike pattern-based methods of cladistic biogeography, it

allows non-hierarchical area relationships. This method has been widely

implemented to evaluate dispersal-vicariance events (Donato et al. 2003;

Miranda-Esquivel 2001; Voelker 2002; Xiang & Soltis 2001; Zink et al. 2000),

and to reconstruct the ancestral area (Anderson 2000; Burns et al. 2002; Davis

et al. 2002; Långtröm 2002).

The purpose of this paper is to analyse distributional patterns of the subgenera

of Simulium within the framework of historical biogeography event-based

methods, by applying dispersal-vicariance analysis (Ronquist 1997). Here an

attempt is made to discern the ancestral areas of the Gondwanian components

of the genus Simulium. This is a preliminary approach and a more refined

analysis, conducted at the species-group level, would ultimately be more

informative.

METHODS

In order to reconstruct the ancestral distributions of the subgenera of Simulium,

the program DIVA 1.2 (Ronquist 1996), is used and a heuristic search using

dispersal-vicariance optimisation is applied (Ronquist 1997). DIVA infers the

ancestral distribution for a taxon and calculates the frequencies of vicariance

and dispersal events among and between the areas under consideration. In

order to do so, the program constructs a three-dimensional cost matrix derived

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from a simple biogeographic model and the phylogenetic information encoded

on the taxon cladogram (Ronquist 1997). Although some other pattern

approaches could be used: treefitter (Ronquist 2001, 2002) and Secondary BPA

(Brooks 1990), DIVA requires the least number of assumptions (contra Weller &

Brooks 2001). The distribution pattern of Simulium subgenera is analysed using

the phylogeny proposed by Miranda-Esquivel & Coscarón (submitted) (Figure

2). The original cladogram is fully resolved, although that illustrated in Figure 2

has some polytomies due to uncertainty in some parts of the cladogram. The

current analysis was conducted using subgenera as terminals. Alternative

topologies found in Miranda-Esquivel & Coscarón (submitted), using different

concavity values in PIWE (vide Goloboff 1993), are also analysed. For the

subgenera S. (Gomphostilbia) Enderlein, 1921, S. (Nevermannia) Enderlein,

1921, and S. (Morops) Enderlein, 1930, the analysis of Otsuka et al. (2001) is

also used (Figure 3).

In total, 13 areas are included, and in the neotropics seven main provinces of

endemism are considered, all defined in previous biogeographic publications

(Coscarón & Coscarón-Arias 1995). For Africa the classical Wallacian sub-

regions are used (Brown & Lomolino 1998). For the sake of this analysis, the

Nearctic and Palaearctic Regions are included and the Australasian and

Oriental Regions are considered as a single biogeographic region. This is not a

novel approach, the combined Australian-Oriental Region, for example, was

used by Ward (2001).

The list of defined areas used in this paper is as follows: (1). Subantarctic +

Central Chile [= Neantartic (= Neantarctica [sic!]) (Monros 1958)], (2). Patagonia

+ Pampa + Monte, (3). Desert + Puna, (4). SE Brasil Mountains + Cerrado, (5).

Guyana + Amazonia, (6). Yungas + North Andean Sub-region, (7). Caribbean +

Pacifica + Mesoamerican Mountains, (8). Nearctic, (9). South Africa, (10).

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Africa: Woodland + Savanna + Desert, (11). Africa: Central Rainforest, (12).

Palaearctic, (13). Australian / Oriental Region.

To infer the age of a clade, the age of vicariance events, as derived from

geological data, is used following Cracraft (2001). To estimate the divergence

time of clades the data from the ‘plates project’ was used (Anonymous 2002).

While other methods to establish divergence time exist (e.g. Huelselbeck &

Imennov 2002), the molecular database for Simulium is currently insufficient to

attempt such analyses.

RESULTS & DISCUSSION

The solutions calculated here require 65 or 66 dispersals and 45 speciation

events, suggesting that current distribution patterns are highly influenced by

dispersal events. Most dispersal events were intra-continental, whereas

vicariance explains most of the inter-continental patterns. Analyses with the

same phylogenetic topologies, but with the biogeographical zones reduced to a

continental scale (Neotropical, Afrotropical, Nearctic, Palaearctic, and the

Australian-Oriental Regions), yielded only 12 dispersal events. These dispersal

events could further be an artefact of uncertainty in the phylogeny, given

different optimisations (vide Figure 4). Dispersal is a critical process in the

cladogenesis of most taxa (Zink et al. 2000), and has also played a critical rôle

in the biogeography of Simulium spp. (Craig et al. 2001; Miranda-Esquivel

2001), the results here, however, suggest that vicariance is the most important

cause of divergence at the subgeneric level.

Austrosimulium Tonnoir, 1925 + Paraustrosimulium Wygodzinsky & Coscarón,

1962, represent the most basal clade in the tribe Simuliini. These two genera

were considered to be sister-groups by Wygodzinsky & Coscarón (1962).

5

Dumbleton (1963) suggested that Austrosimulium was derived from Gigantodax

Enderlein, 1925, and that they both originated in South America or in Antarctica.

Crosskey (1987, and more recently) did not include both taxa in the same clade.

According to the present analysis, the ancestral area for Austrosimulium +

Paraustrosimulium is Australia + Subantarctic province, indicating a clear

Gondwanian relationship with a southern ocean baseline (Craw et al. 1999).

Many southern South America/Australia events have been dated as late

Cretaceous, because the isolation between Australia and southern South

America began with opening of the Drake Passage about 36 Mya (Pascual

1998), and this isolation increased with the formation of ice caps on Antarctica

(Crosskey 1990). Placing the origin of the ice caps at the middle Miocene (13-

10 Mya), the basal position of the clade must be prior to this date, and could be

as early as the opening of the South Atlantic circa 135 Mya. In agreement with

Crosskey (1990), the taxon inhabited Antarctica. Lutzsimulium hirticosta Lutz,

1909, could be the sister-group to the previous clade. This sister-group

relationship reinforces the idea of a Gondwanian relationship and does not

influence the estimated divergence time.

This analysis found that the Central Rainforest + Savanna was the ancestral

area for the afrotropical clade: S. (Lewisellum) Crosskey, 1969 + S.

(Phoretomyia) Crosskey, 1969. This clade is recovered in all the phylogenetic

analyses. It is a young clade that may be dated as early as the post Cretaceous

or Tertiary. The sister-group to this taxon may be S. (Psilozia) vittatum

Enderlein, 1936, as suggested by some of these analyses, but this proposed

relationship infers an unlikely vicariance or dispersal event. For the clade S.

(Pomeroyellum) Rubtsov, 1962 + S. (Meilloniellum) Rubtsov, 1962 + S.

(Nevermannia) ruficorne-group, the ancestral area is also the Central Rainforest

+ Savanna, and this is also a recent divergence. African and South American

Prosimuliini (sensu Coscarón 1991), may also have had a Godwanian

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distribution. Neotropical Araucnephia Wygodzinsky & Coscarón, 1973 and

Araucnephiodes Wygodzinsky & Coscarón, 1973, are phenotypically more

similar to the South African Paracnephia (Paracnephia) Rubtsov, 1962, than to

any other neotropical genus (Coscarón & Coscarón-Arias 2002), and

Paracnephia has a clear Gondwanian origin (Crosskey & Howard 1997).

The ancestral area for the Australian-oriental clade, S. (Inseliellum), S.

(Morops), S. (Gomphostilbia), and S. (Nevermannia) loutetense-group, is the

Australian-Oriental Region + Africa. This could be a recent divergence. Craig et

al. (2001) considered the divergence for S. (Gomphostilbia), S. (Inseliellum), S.

(Morops) and S. (Wallacellum) Takaoka, 1983, to be 20 Mya as a dispersal

across SE Asia; while S. (Hebridosimulium) Grenier & Rageau, 1961, dispersed

to Fiji and Vanuatu circa 10-6 Mya. Otsuka et al. (2001), applied molecular data

to analyse the Australian-oriental clade (Figure 3), but did not include S.

(Inseliellum), S. (Hebridosimulium) and S. (Wallacellum). This finding suggests

that S. (Nevermannia) feuerborni-group could have a palaearctic or oriental

origin. The divergence of the Australian-oriental clade that includes both

subgroups (Africa + Australian-oriental clades) may be placed in the lower

Cretaceous. This pattern, with an Indian Ocean baseline (Craw et al. 1999),

joining the Africa + Australian-Oriental Regions has been found in other

analyses. For example, Ward (2001) found an African origin for the genus

Tetraponera Smith, 1852, and four independent origins for the Australian-

oriental clades. He placed the divergence time for Tetraponera in the

Cretaceous. Holloway & Hall (1998) showed a similar pattern with some groups

of Lepidoptera (Troidini, Callidulidae, Uraniidae and Castniidae), although there

was no common pattern for the four taxa.

Neotropical Simulium subgenera are divided into three subclades. The first

clade, S. (Ectemnaspis) Enderlein, 1934, S. (Psilopelmia) Enderlein, 1934, and

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S. (Chirostilbia) Enderlein, 1921, represents a well-supported node (Coscarón

1987), for which the ancestral area was found to be the entire Neotropical Sub-

region, excluding Deserts and Puna. Coscarón et al. (1996) analysis suggests

that the Nearctic component of the S. (Psilopelmia) clade might be a basal

divergence in the subgenus and that this divergence within the genus is rather

recent, though they did not suggest a divergence time. The sister-group to S.

(Ectemnaspis), S. (Psilopelmia), and S. (Chirostilbia) is S. (Psaroniocompsa)

sensu lato Enderlein, 1934, S. (Inaequalium) Coscarón & Wygodzinsky, 1984,

S. (Aspathia) Enderlein, 1935, and S. (Notolepria) Enderlein, 1930. This

research supports the hypothesis that the latter clade is neotropical, with

Mesoamerica (Caribbean + Pacifica + Mesoamerican Mountains) as the

ancestral area. It is also a recent divergence.

In the neotropical realm S. blancasi-group + S. (Pternaspatha) Enderlein, 1930,

form a well-established clade (Coscarón 1987; Coscarón & Coscarón-Arias

1996; Miranda-Esquivel & Coscarón submitted). Its ancestral area is

Desert/Puna. If S. (Afrosimulium) Crosskey, 1969, is considered as the sister-

group to this taxon, as suggested in some of these optimisations (Figure 4), the

ancestral area for this clade (Afrosimulium + (blancasi-group + Pternaspatha))

is South Africa - Desert/Puna. This indicates an ancient origin for the clade in

the early Cretaceous (circa 110 Mya) or even late Jurassic, before the

separation of southern Gondwana and the formation of the southern Atlantic

Ocean (Pitman et al. 1993). The weather in the middle America-south and

central Africa from the Volgian onwards, became a winter-wet zone, after a

longer desert zone period during the Jurassic (Rees et al. 2000). This shall

undoubtedly be regarded as a rather controversial date, as the oldest known

fossil for the family is a Prosimulium-like genus that has been dated as

Jurassic/Cretaceous, and because Prosimuliini sensu Currie (1988) is older

than Simuliini. A suggested solution to solve the too early date is a dispersal

8

event across a narrow proto-Atlantic. This solution could be possible but no

longer than 100 Mya. There is no evidence of extant species belonging to the

clade (Afrosimulium + (blancasi-group + Pternaspatha)) in the southern Atlantic

islands. The alternative solution, found with other optimisations, is for S.

(Afrosimulium) to be placed as the sister-group of S. (Methomphalus) Enderlein,

1935, and the ancestral area for this taxon (Afrosimulium + Methomphalus)

would be South Africa + Savanna.

The clade S. (Thyrsopelma) Enderlein, 1934, S. (Trichodagmia) Enderlein,

1934, S. (Freemanellum) Crosskey, 1969, S. (Anasolen) Enderlein, 1930, S.

(Xenosimulium) Crosskey, 1969 and S. (Hemicnetha) Enderlein, 1934, is well

supported (Miranda-Esquivel & Coscarón 2001), which falsifies the hypothesis

that all neotropical Simulium spp. form a monophyletic clade (Coscarón 1987).

These results show that the ancestral area of S. (Thyrsopelma) + S.

(Trichodagmia) is Amazonia sensu latissimo (Guyana + Amazonia + Cerrado +

SE Brasil), while the ancestral area for the whole clade (Thyrsopelma +

Trichodagmia + Freemanellum + Anasolen + Xenosimulium +Hemicnetha) is

Amazonia s. l. + Africa. All subgenera are found exclusively in Africa or South

America, suggesting that the vicariance event separating them was in the early

Cretaceous, circa 100 Mya, or as early as the beginning of the Cretaceous

(Cracraft 2001; Anonymous 2002). Since the late Jurassic the weather in the

Amazonia-Africa junction has been summer-wet (Rees et al. 2000). The African

subgenera S. (Methomphalus) together with S. (Edwarsellum) Enderlein, 1921,

and S. (Byssodon) griseicolle-group are basal to the Gondwanian clade (vide

Figure 2). This result could infer a Gondwanian origin to these subgenera.

According to these results, the ancestral area of S. (Edwarsellum) + S.

(Byssodon) griseicolle-group could be Africa or the palaearctic. It is therefore

difficult to explain the distribution of S. (Byssodon) Enderlein, 1925. It could be a

palaearctic subgenus with a broad dispersal, or a non-monophyletic taxon such

9

as S. (Nevermannia), where various components are found in different regions.

The neotropical, Simulium oviedoi species-group is a problematic taxon. Some

of these results suggest a sister-group relationship with S. (Ectemnaspis) + S.

(Psilopelmia) + S. (Chirostilbia) (Coscarón 1987), or alternately a sister-group

with Hemicnetha (Ramírez-Pérez 1971). The affinities are still unclear, but

neither of the two possible solutions alter the optimisation of the ancestral area.

The nearctic subgenus S. (Psilozia) is placed as a basal taxon in this analysis,

and this may indicate a ‘remote’ relationship to the nearctic/palaearctic

Simulium fauna. This hypothesis could be tested only with a phylogenetic

analysis that includes all nearctic and palaearctic subgenera.

The Neotropical and the Afrotropical Regions are complex areas that exhibit

different affinities (Figure 5). The Neotropical Sub-region is closest in simuliid

phylogenetic similarity to central Africa [the Atlantic Ocean baseline, Craw et al.

(1999)]. This pattern is evident in the S. (Thyrsopelma), S. (Trichodagmia), S.

(Freemanellum), S. (Anasolen) and S. (Xenosimulium) clade. The Andean Sub-

region is related to the Australian-Oriental Region [the southern Ocean

baseline, Craw et al. (1999)]. This pattern is present in the divergence between

the sister taxa Austrosimulium and Paraustrosimulium. Africa is also related to

the Australian–Oriental Region [the Indian Ocean baseline, Craw et al. (1999)].

This pattern is evident in the divergence of the sister taxa S. (Inseliellum), S.

(Morops), S. (Gomphostilbia), and S. (Nevermannia) loutetense, where the

basal African taxon S. (Nevermannia) loutetense is related to the Australian-

oriental clade S. (Inseliellum), S. (Morops), and S. (Gomphostilbia).

The area cladogram presented here, ((Australian-Oriental, Southern South

America) + ((Africa, Australian-Oriental) + ((Africa, Amazonia), South

America))), do not completely agree with the general pattern proposed for

10

Gondwanian clades (Linder & Crisp 1995). Although, some sister areas do

agree, such as Africa + Oriental, and Southern South America + Australia. The

only difference lies in the basal divergence between Southern South America +

Australia. The interpretation is that this divergence is dated towards the time of

the opening of the South Atlantic 135 Mya, suggesting a connection via

Antarctica. This hypothesis is different than that proposed by Crosskey (1990)

and it is in better agreement with the general area cladogram proposed by

Linder & Crisp (1995). Nonetheless, the divergence from the general pattern

may also be assigned to dispersal. These results are not unique in differing

from the general pattern, incongruence has been found with other groups (e.g.

Edgecombe et al. 2002).

Not surprisingly, the ancestral area for the whole clade under analysis is a zone

that includes all the areas considered. Such a wide distribution means that

Simulium is older than the break-up of the plates considered here, as old as the

middle Jurassic (circa 160 Mya), when the Tethys Sea divided Pangea into

Laurasia and Gondwana. This is older than the earliest fossil Kovalevimyia. If

‘Simuliini’ is more recent than the basal Simulium clade, then the origin of the

family must predate the late Jurassic. Parasimuliinae and Prosimuliini (sensu

Currie 1988) are the most basal nodes in the family (Moulton 2000), and both

taxa are holarctic. A possible interpretation to that topology is that the ancestral

area for the family is either the holarctic as Currie (1988) proposed for

Prosimuliini, or Pangea. Depending on the optimisation Simulium may have had

a Pangean distribution. This distribution reinforces the idea that the family has

an ancient origin, perhaps early Jurassic, which was the lowest boundary

proposed by Crosskey (1990). These findings open questions about the

divergence time and the geographical origin for Simulium, and more data, both

molecular and morphological, are necessary to attempt a more refined answer.

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ACKNOWLEDGEMENTS

We are grateful to Mike Sharkey (University of Kentuchy, USA), Mariano

Donato (Museo de la Plata, Argentina), Paula Posadas (Museo Egidio Feruglio,

Argentina), Ashley Kirk-Spriggs (National Museum of Namibia, Windhoek) and

two anonymous reviewers for valuable comments on the manuscript. Nelida

Caligaris prepared the illustrations. This research was supported in part by the

grant DIF-Ciencias-UIS 5112. The first author is indebted to Colciencias

(Consejo Nacional de Ciencia y Técnica, Colombia) and to DIF-Ciencias-UIS for

their financial support.

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[Legends to Figures]

Figure 1. Area cladogram illustrating biogeographical patterns in the Neotropical

Region utilising subgenera of Simulium Latreille (after Miranda-Esquivel 2001).

Figure 2. Conservative cladogram illustrating the main cladogenetic events for

subgenera of Simulium Latreille (after Miranda-Esquivel & Coscarón

(submitted)). Taxa abbreviations: B. = Byssodon; N. = Nevermannia; S. =

Simulium.

Figure 3. Cladogram after Otsuka et al. (2001). This is an alternative solution to

the Austral-oriental Taxa abbreviation: S. = Simulium.

Figure 4. Simulium Latreille subgenera cladogram using a concavity value of

one or two. This is an alternative solution to Figure 2 showing the clade

Afrosimulium + S. blancasi-group + S. (Pternaspatha) as a basal node to

Simulium clade. Taxa abbreviations: B. = Byssodon; N. = Nevermannia; S. =

Simulium.

Figure 5. Distributional patterns for neotropical, afrotropical and Australian-

oriental Simulium Latreille subgenera. Some additional genera have been

included to illustrate general affinities [after Crosskey (1990). Dates from

Cracraft (2001) and the ‘Plates project’ (Anonymous 2002)].

21

Mesoamerica

Nearctic

Pacifica

Caribbean

Yungas

North Andes

Desert

Puna

Patagonia

Monte

Subantarctic

Central Chile

SE Brasil

Guyana

Cerrado

Amazonia

Cnesia dissimilis Lutzsimulium simplicicolor-group Lutzsimulium hirticosta-group Paraustrosimulium Austrosimulium Simulium (Psilozia) vittatum S. (Afrosimulium) S. (Phoretomyia) S. (Lewisellum) S. (N.) ruficorne-group S. (Meilloniellum) S. (Pomeroyellum)

S. (N.) loutetense-group S. (Morops) S. (Inseliellum) S. (Gomphostilbia) S. blancasi-group S. (Pternaspatha) S. (Chirostilbia) S. (Ectemnaspis) S. (Psilopelmia) S. (Notolepria) S. (Psaroniocompsa) S. (Inaequalium) S. (Aspathia) S. (B.) griseicolle-group S. (Edwarsellum) S. (Methomphalus) S. oviedoi-group S. (Hearlea) S. (Hemicnetha) S. (Anasolen) S. (Freemaniellum) S. (Xenosimulium) S. (Trichodagmia) S. (Thyrsopelma)

African clade

Australia-oriental clade

Neotropical clade

Gondwanian clade

Prosimulium

S. (N.) ruficorne group

S. (Morops)

S. (N.) feuerborni group

S. (N.) vernum group

S. (Gomphostilbia)

S. konoi

S. (Simulium)

Cnesia dissimilis Lutzsimulium simplicicolor-group Lutzsimulium hirticosta-group Paraustrosimulium Austrosimulium S. (Afrosimulium) S. blancasi-group S. (Pternaspatha) S. (Edwarsellum) S. (Psilozia) vittatum S. (Phoretomyia) S. (Lewisellum) S. (Methomphalus) S. (Thyrsopelma) S. (Trichodagmia) S. (Xenosimulium) S. (Freemaniellum) S. (Anasolen) S. (Hearlea) S. (Hemicnetha) S. (Chirostilbia) S. oviedoi-group S. (Psilopelmia) S. (Ectemnaspis) S. (Notolepria) S. (Aspathia) S. (Inaequalium) S. (Psaroniocompsa) S. (B.) griseicolle-group S. (Pomeroyellum) S. (Meilloniellum) S. (N.) ruficorne-group S. (N.) loutetense-group S. (Gomphostilbia) S. (Inseliellum) S. (Morops)

1