Table Of ContentZoological Journal of the Linnean Society, 2017, 181, 151–188. With 38 figures.
Phylogeny of Phorinae sensu latu (Diptera: Phoridae)
inferred from a morphological analysis with
comprehensive taxon sampling and an uncommon
method of character coding
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DANILO CÉSAR AMENT* w
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Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de e
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São Paulo, Avenida Bandeirantes 3900, 14040–901 Ribeirão Preto, SP, Brazil fro
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Received 27 June 2016; revised 11 October 2016; accepted for publication 9 November 2016 ttp
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Few comprehensive works have dealt with supra-generic relationships of the Phoridae. These resulted in different m
classifications, especially regarding the composition and internal relationships of the subfamily Phorinae. Although ic.o
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a consensus regarding Phorinae delimitation seems to have been recently achieved, relationships among the genera p
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in this subfamily are still contentious. The present analysis intended to recover the relationships between Phorinae o
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genera recognizing well-supported groups within this subfamily. A strategy distinct to those previously applied to /z
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this problem was conducted. Most characters proposed in the literature were verified and included – even those o
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previously considered ‘highly homoplastic’. Characters were re-delimited or re-coded when necessary and new char- n
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acters were also proposed. Characters were coded according to hierarchical hypotheses of homology in cases of special a
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state-spaces. Taxonomic sampling was designed to represent as best as possible the morphological diversity of the /a
characters chosen and to reduce possible long branches. This included the first inclusion of 34 fossil Phorinae taxa in rticle
a phylogenetic analysis of the subfamily. The matrix consisted of 125 characters from 141 taxa and resulted in three /1
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most-parsimonious trees. The strict consensus of these trees corroborated several large groups that had been previ- 1/1
ously proposed in similar or equal compositions, including Euphorida, Neophorida, Phorinae s.l., Bothroprosopa- /1
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group, Thaumatoxena-group and Hypocera-subgroup. It also revealed several new well-supported small groups of 1/3
genera and large clades. The results showed that there is indeed relevant phylogenetic information in characters 07
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previously thought to be highly homoplastic, and that these characters can be optimized with reasonable congruence 1
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when analysed with thorough taxonomic sampling. 8 b
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ADDITIONAL KEYWORDS: classification – morphology – phorids – phylogenetic analysis. u
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INTRODUCTION example, estimates indicate that only one-tenth of spe- ug
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Phoridae is a family of small cyclorrhaphan flies – 0.4–6 cies in certain genera are known (Brown, 2004). st 2
There are three main works regarding the supra- 0
mm in body length – with about 4000 described spe- generic classification of the Phoridae: Schmitz (1929), 21
cies. They are distributed worldwide and, in general,
Brown (1992) and Brown, Amorim & Kung (2015).
can be easily recognized by their intermittent locomo-
Schmitz (1929) divided the family into four subfami-
tion, reduced wing venation and longitudinal setal pali-
lies such as Aenigmatiinae, Metopininae, Phorinae
sades, at least on the hind tarsomere. The family has an
and Thaumatoxeninae and created the family
astounding diversity of larval habits, containing preda-
Termitoxeniidae for the genera now considered the
tors, parasites, parasitoids, herbivorous, saprophagous
termitophilous phorids. He delimited the Phorinae
and fungivorous species (Disney, 1994). However, many
in a relatively wide sense, defining them as winged
aspects of the Phoridae are still poorly known. For
phorids with upward pointing supra-antennal setae,
tibiae almost always with bristles and anepisternum
*E-mail: [email protected] furrow generally absent.
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188 151
152 D. C. AMENT
Brown’s (1992) phylogenetic analysis focused on retrieve characters information at sufficient levels of
the Nearctic genera of the family and resulted in a the phylogeny, a special investment in taxon sampling
classification conflicting with that of Schmitz, espe- was made to include as best as possible the generic
cially regarding the subfamily Phorinae. According to diversity regarding the characters and to shorten
Brown’s phylogeny, the Phorinae (sensu Schmitz) are the evolutionary distance between the taxa sampled.
paraphyletic, justifying their division into three sub- Additionally, uncommon coding methods using step-
families: Conicerinae, Hypocerinae and Phorinae s.s. matrices were also applied to recover different levels
Brown’s re-delimitation was opposed by Disney (1993), of phylogenetic information in certain characters.
and a debate on homology and other aspects of phy-
logenetic methodology followed (Disney, 1994, 1995,
1996; Brown, 1995a). In spite of the disagreements, no
MATERIAL AND METHODS D
new classification was proposed until recently. o
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Brown et al. (2015) performed a phylogenetic analy- Specimens of 141 species of 90 genera, housed mostly n
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sis of the family with an extensive sampling of gen- in the Natural History Museum of Los Angeles a
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era and the inclusion of 37 new characters based on County (LACM) and the Museu de Zoologia de São d
thorax morphology. They recovered Phorinae s.s. as Paulo (MZSP), were analysed (Table S1). Most speci- fro
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paraphyletic and re-delimited the subfamily as com- mens were dried by the curatorial team of the LACM h
prising all phorids not included in the Sciadocerinae, with hexamethyldisilazane (Brown, 1993a) and were ttp
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Chonocephalinae, Metopininae or Termitoxeniinae. in excellent condition. Specimens of almost all spe- ://a
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In spite of the advances made in the delimitation of cies had wing and posterior leg permanently slide- a
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subfamilies, the authors considered the phylogenetic mounted. Certain structures were selected for more em
resolution within the Phorinae doubtful and deserving detailed study, such as the antennae and palpi, and ic
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of more investigation. were dissected and prepared in temporary slides with up
Relationships between phorine genera remain glycerin for observation under a compound microscope. .co
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a controversial topic mostly because of the rapid, Hypopygia were dissected and cleared in lactic /z
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homoplastic evolution of some of the most important acid for a better preservation of membranous parts o
and well-delimited characters within the subfamily (Burington, 2010). Afterwards, they were temporarily linn
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(Disney, 1991; Brown, 1992). Whether these characters mounted on slides with triangular excavations filled a
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sohdooulolgdi cbael isntcrlautdeegdie isn b tehset arencaolvyesirs tahnedir winhfiocrhm maetitohn- wofi tahlm gloyscte arliln s fpoerc iilelus ssttruadtiieodn w(Aemree niltlu, 2s0tr1a5t)e. dH iynp dooprysgaila, /article
at the appropriate phylogenetic level are questions ventral and both left and right lateral views following /1
8
1
which were approached differently by Brown (1992) the taxonomic revisions of Ament & Amorim (2013) /1
and Brown et al. (2015). and Ament (2012, 2014). Female terminalia, which /15
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Brown (1992) used genera and subfamilies as ter- frequently have segments 7–10 internally telescoped, /3
0
minal taxa inferring their states through a Bauplan were cleared in KOH for better dissolution of their mus- 7
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method. However, in terms of the characters analysed, culature facilitating the extrusion of these segments. 12
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this lacked a large portion of the diversity of the supra- Along with the study of complete specimens, close-up b
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specific taxa now known to exist. In contrast, Brown et illustrations were also prepared of certain structures g
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al. (2015) used the exemplar method, placing species for almost all taxa and grouped in pictorial matrices es
as terminal taxa. Although the authors stated that to facilitate the morphological comparison and identi- t o
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they examined the genera for intra-generic variation, fication of characters. These include the head (dorsal 1
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they sampled only one species per genus and did not and lateral views), wing, posterior femur, fore, mid and A
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consider polymorphism in the matrix. Therefore, it hind tibia, apex of wing vein R , R and wing base gu
4 + 5 1 s
seems that the Brown et al. (2015) strategy of analy- (plates in Figs S8–S39). Whenever possible, the collec- t 2
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sis was not to include rapidly evolving characters or to tion numbers of illustrated specimens were indicated. 2
1
use them with a restricted coding, such as character 59 Fossils and other undescribed materials were referred
‘Male first flagellomere in sciadocerines’. to by their LACM identification labels. Fossils, in par-
The aim of the present analysis was to recover the ticular, were not illustrated as they will be described in
relationships between Phorinae genera and recog- future by Dr Brown.
nize well-supported groups within the subfamily. To Terminology follows Cumming & Wood (2009).
reach this objective, I used a strategy distinct from Hypopygial plates were referenced according to Brown
those previously applied to this problem. It differed in (1992), with some exceptions as discussed in the char-
that it tried to recover the phylogenetic information acters. Parts of the female terminalia were named
of most well-delimited characters proposed for the according to Nakayama (2007). The group name
family, including characters known to be homoplastic Euphorida refers to Termitoxeniinae + Metopininae +
at certain levels, using species as terminal taxa. To Phorinae, as in Brown (2007), and Neophorida refers
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
PHYLOGENY OF PHORINAE 153
to Metopininae + Phorinae, as in Brown et al. (2015). Hypotheses of homology at different levels are generally
The names Hypocera-subgroup and Bothroprosopa- coded through a step-matrix of differential transition
group refer to groupings originally proposed by Brown costs between states, a character state tree (Farris, Kluge
(1992), although the latter was recovered here in a & Eckhardt, 1970), or as contingent characters (Forey
slightly different composition. & Kitching, 2000). These include ‘additive’ coding and
strategies that use other types of step-matrices (Sankoff
& Rousseau, 1975). Non-hierarchical coding is usually
represented by multistate non-additive characters.
PHYLOGENETIC ANALYSIS
In essence, there is no difference between combin-
Taxon sampling ing different morphological conditions in a single char-
Species of 52 of the 60 phorine genera were included. acter state and combining character states in a more
D
Most of the genera not included belong to the inclusive homology hypothesis. Both follow the same ow
Aenigmatiinae sensu Brown (1992) and are known only criteria (topographic, connection by intermediates, nlo
from females. Species sampling followed the divergent lin- special similarity) and are interpreted according to ad
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epaagreasp hmyeltehtiocd o (rY heaigtehsl,y 1 d9i9v5e)r,s we irtehg gaerndeinrag stuhsep cehctaerda cttoe bres t1h9e9 2a)u. xFiolilalorwy ipnrgi nthciipsl leo gofi cH, Ie nconnigs i(d1e9r6e6d) h(iWerialkricnhsiocanl, d from
used being sampled more extensively. Representatives of coding strategies valid for recovering homology infor- h
all other Phoridae subfamilies were sampled following mation and used additive and other types of step- ttps
the same criteria and included as outgroups. matrices to code appropriate characters (examples in ://a
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Fossil phorine were included in a phylogenetic analy- Fig. 1). Homology hypotheses of different levels can ad
sis of the subfamily for the first time. According to the be incorporated into the analysis through the scor- em
simulations by Huelsenbeck (1991), fossils are espe- ing of differential transition costs between states. A ic.o
cially important in analyses of rapidly evolving charac- transition between states with a lower cost will favour up
ters and they represent lineages close to the phylogeny the homology between these states. As the transition .co
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nodes or across long time intervals. Indeed, this seems costs between these states are still a positive valour, /z
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tooth beer trheae scoansse, mwoitsht othf eth per aesmebnetr a snpaelcyiseiss ,i nacsl,u admedo ndgo lweistsh iinnc tlhuissi vme ohroem inoclolugsyi vhey phoytphoetsheess iasr.e also recovered olinn
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not fit into any extant genera. In total, 34 Baltic or Six measurement-based characters were included an
oonldee ‘rp raimorbiperh osrpiencei’e as nwde 3re2 iPnhcoluridneade ( so.nl.e). Sciadocerinae, aans dc toenstteindu uonudse rc thwaor adicftfeerresn, ta ccocdoirndgi mnge tthoo dGs.o Floirbsotflfy,, /article
Mattoni & Quinteros (2006), and secondly, as discrete /1
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CharaCTer sampling astdadteitsi.v Beo cthh athraecstee mrse twhiotdhs rraesnugletesd d iinv itdheed s ainmteo tnhirneee /1/15
Most characters previously proposed for Phoridae phy- most parsimonious trees (MPTs) but with slightly 1/3
logeny were included, although sometimes using a dif- higher support values using the second. These char- 07
5
ferent coding. These were re-checked for all taxa and acters were therefore converted into discrete variables 12
their states were frequently scored differently to the for analysis in this paper. 8 b
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original work. New characters, mostly from the mor- g
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phology of wing sensilla, tibial chaetotaxy, hypopyg- e
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ium and oviscape, were also added. Most characters WeighTing t on
were constructed based on observation of males, with 1
Some strategies of coding hierarchical hypotheses of 0
many being exclusive to them. Characters exclusive to A
homology applied here require the assignment of dif- u
females are indicated as such. g
ferential transition costs between states. Consequently, us
characters coded in this way necessarily have at least t 20
2
CharaCTer Coding one transition cost higher or lower than the transition 1
costs of binary and multistate non-additive characters.
The state space of a morphological character usually
To balance out these differences, the maximum transi-
comprises a wide diversity of conditions of different mor-
tion cost of most additive characters was made equal to
phological disparities that are grouped as a few charac-
the transition costs of other characters (stipulated as 6).
ter states (Pogue & Mickevich, 1990). Such state spaces
often contain phylogenetic information of homologies
of different levels of inclusiveness. Two types of coding
strategies are frequently used in these cases: those that sofTWare
take into consideration hypotheses of homology at differ- The matrix was built in Winclada (Nixon, 1999) and
ent levels (hierarchical) and those that recover hypoth- analysed in TNT (Goloboff, Farris & Nixon, 2008).
eses of homology at only one level (non-hierarchical). Bremer support values were calculated in TNT.
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
154 D. C. AMENT
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Figure 1. Transition costs and their resultant homologies (represented as dashed forms) of two hierarchical coding n
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schemes: an additive multistate character (left) and a multistate character with another type of step-matrix (right). a
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RESULTS hypothesizing relationships between groups external from
to the Thaumatoxena-group (topic 2 of discussion). h
The matrix consisted of 125 characters for 141 taxa ttp
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(Table S2). Of these characters, 75 are binary, 13 ://a
multistate non-additive and 38 multistate with dif- CharaCTers ca
d
ferential transition costs between states – the latter e
Several of the characters included in this study were m
being divided into 26 coded as discrete additive and ic
previously proposed in the literature. For some of these, .o
12 with other types of step-matrices. The analysis of u
the coding used was substantially different from that p
this matrix resulted in three MPTs with a consistency .c
originally proposed. These are discussed in the light of o
index of 22 and retention index of 64 (mean of all char- m
the coding methods used and their implications. Many /z
acters except for the six measurement characters). The o
strict consensus of these trees, which is equal to one other characters proposed in the literature in dis- olin
tinct contexts (e.g. Schmitz, 1929, 1951) or without an n
MPT, is used to discuss the phylogenetic results (Fig. ea
explicit coding method (e.g. Disney, 1986, 1991, 2003) n
2). /a
Several of the large groups recovered are consist- were not discussed given the incomparability of their rtic
coding methods with the present analysis. Most com- le
ent with Brown et al. (2015), namely the Phoridae, /1
parisons are made with the works of Brown (1992) and 8
Neophorida (Metopininae + Phorinae), Metopininae 1
Brown et al. (2015), which incorporated most hypoth- /1
and its subgroups, the Phorinae and the Hypocera- /1
eses of these previous studies. 5
subgroup. The sister-group relationship between Possible correlations between characters were con- 1/3
Chonocephalinae and Termitoxeniinae differs from 07
sidered and strategies were taken to avoid possible 5
the hypothesis Chonocephalinae + (Neophorida + 1
biases in the analysis. The transition costs used to 2
Termitoxeniinae) in Brown et al. (2015) (topic 1 of recover more inclusive hypotheses of homology are 8 by
discussion). Conicerinae sensu Brown (1992) was also indicated. A simplified representation of each char- gu
recovered here as polyphyletic, with only Contopteryx e
acter optimization in the strict consensus is given to s
Schmitz and Darwiniphora Schmitz shown as closely t o
show their structure in the phylogeny. Consistency n
related to the type genus Conicera Meigen. The close 1
and retention indices are also used to discuss charac- 0
relationship between Aenigmatiinae sensu Brown A
ter evolution. u
(1992), Neopleurophora Brown, Chaetocnemistoptera g
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Borgmeier, Hirotophora Brown, Kung & Amorim and t 2
the ‘hypocerines’ is recovered as in Brown et al. (2015), measuremenT CharaCTers 02
1
but with considerable differences in their relative posi- (disCreTized)
tions (topic 3 of discussion). The group formed by these
1. Frontal height/width (Fig. S1, optimization; Fig. 3,
taxa is hence named the Thaumatoxena-group.
states discrimination).
Within the Thaumatoxena-group, the Hypocera-
group of Brown (1992), also referred to as the ‘hypocer- Original interval: 0.3–4.6 (scaled to states 0–9).
ines’ in Brown et al. (2015), also form a clade, here named C.I. = 15; R.I. = 35.
the Hypocera-subgroup. Most remaining Hypocerinae In spite of the considerable morphological varia-
sensu Brown (1992), or the ‘Bothroprosopa-group’, are tion of frontal dimensions among Phoridae genera,
recovered here as a distinct group including genera this information has never been coded in a charac-
not previously considered closely related. The present ter in a comprehensive phylogenetic analysis of the
phylogeny also differs from Brown et al.’s (2015) by family. Brown (1992) inferred ‘reduced frons’ as a
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
PHYLOGENY OF PHORINAE 155
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Figure 2. Strict consensus of the three most parsimonious trees obtained. Bremer support values are indicated under each
node. Extinct taxa in brown.
synapomorphy of Conicerinae, but did not include this the present character as the ratio frontal height/width
information in any character. Brown et al. (2015), in dividing this ratio values into nine states and treating
character 54 ‘Male frons: (0) narrowed, (1) broad like them as additive.
that of female’, attributed the state of narrowed frons Reduction in frontal height is possibly correlated
only to Ironomyiidae and more external groups. To with reduction in number of frontal setae in some taxa
recover the different levels of phylogenetic information as Conicera and Contopteryx. However, there are taxa
of the frontal dimensions, I coded this information in with broad frons with no frontal setae and taxa with
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
156 D. C. AMENT
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Figure 3. Examples of different conditions of character 1 (frontal height/width). d fro
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Figure 4. Examples of different conditions of character 2 (hind femur height/length). t o
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narrow frons with three rows of frontal setae, which Original interval: 0.15–0.45 (scaled to states 0–9). A
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are evidence of at least partial independence between C.I. = 13; R.I. = 31. g
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these characters. The development of the first flagel- Brown et al. (2015) coded the shape of the posterior t 2
lomere (character 17) and the frontal fold (character femur as two discrete states in character 26 ‘Hind 02
1
16) of some taxa are also conditions probably related femur and coxa: (0) cylindrical (1) posteriorly flat-
to the frontal dimensions. tened’. I considered the different levels of anteropos-
The increase in frontal height/width is notable at terior flattening of the hind femur best represented
the Euphorida base. Within Euphorida, relatively nar- by a continuous measure related to the height/length
rower frons is independently optimized for the clades ratio of this structure. To recover the different lev-
Hirotophora + Neopleurophora + Chaetocnemistoptera, els of phylogenetic information of this ratio, I discre-
Postoptica Disney + Termitoxeniinae and the grouping tized it in nine states and treated them as an additive
of Phora Latreille species (Fig. S1). character.
A relatively large femur was optimized at the
2. Hind femur height/length (Fig. S1, optimization; Euphorida base with reversions to a relatively thin
Fig. 4, states discrimination). femur in some small groups such as the clade Misotermes
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
PHYLOGENY OF PHORINAE 157
Schmitz + Aenigmatistes Shelford, part of the 4. Foremetatarsus width/length (Fig. S2, optimiza-
Bothroprosopa-group and part of Phora species (Fig. S1). tion; Fig. 6, states discrimination).
Original interval: 0.10–0.58 (scaled to states 0–9).
3. Vein C length/wing length (Fig. S2, optimization;
C.I. = 11; R.I. = 19.
Fig. 5, states discrimination).
Foremetatarsus dimensions were used here for the
Original interval: 0.18–1.0 (scaled to states 0–9). first time in a phylogenetic analysis of the family. This
C.I. = 12; R.I. = 34. character was optimized with a plastic evolution in the
The relative length of the costal vein is used in tree but being informative in less inclusive levels. A rela-
Phoridae taxonomy in distinct contexts from spe- tively short foremetatarsus was recovered as synapomor-
cies descriptions to inferences of relationships. phic for Chonocephalus Wandolleck + Termitoxeniinae
However, the only use of this information as a char- and Euryphora Schmitz + Aenigmatias Meinert. D
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acter of a comprehensive phylogenetic analysis w
5. Scutellum length/width (Fig. S3, optimization; n
was character 2 of table 8 of Brown (1992) ‘Costa lo
exceptionally long’, which was recovered as a syna- Fig. 7, states discrimination). ade
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pomorphy of Ceratusa Borgmeier + Cyphometopis Original interval: 0.22–0.77 (scaled to states 0–9). fro
Borgmeier. This character is coded here with nine C.I. = 12; R.I. = 27. m
h
additive states given its broad and continuous Scutellum dimensions are possibly related to char- ttp
vaAri ashtioornte.ning of vein C was optimized at the base of a(0c)t eclro 3s4e toofg Betrhoewr,n ( 1et) abrl.o (a2d0l1y5 s)e ‘pPaorsatetreido’r, wschuitcehl lwara ss ectoane-: s://ac
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Jealia Brown + Euphorida, while even greater reduc- sidered by these authors as correlated with body size. de
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tions in this vein length are optimized independently Scutellum length/width ratio was chosen among these ic
for small groups within Euphorida (Fig. S2). three measures as it can be measured independently .o
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Figure 5. Examples of different conditions of character 3 (vein C length/wing length). gu
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Figure 6. Examples of different conditions of character 4 (foremetatarsus width/length).
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
158 D. C. AMENT
from other characteristics such as the number and Brown (1992) referred to this character as an example
arrangement of scutellum setae. of possible atavism in the evolution of the family, rec-
A relatively narrow scutellum was optimized in ognizing this furrow in phorids as a possible re-expres-
most groups of Phoridae with two major groups pre- sion of the furrow of other Diptera families. According
senting a wider scutellum, the Metopininae and the to Brown, the furrow in Phora and Coniceromyia
Thaumatoxena-group (Fig. S3). Borgmeier would be independent synapomorphies of
both of these genera. The present analysis corrobo-
6. Distance R –costal apex/C length (Fig. S3, optimi-
1 rated these independent origins optimizing two other
zation; Fig. 8, states discrimination).
presences of this furrow within Metopininae.
Original interval: 0.22–0.59 (scaled to states 0–9). Some genera as Archiphora have a vestigial frontal
C.I. = 19; R.I. = 29. furrow restricted to a concavity at the venter of the
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Besides the relative costal length (character 3), the frons, a condition which was coded here as furrow o
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radial system varies also regarding the R relative absent. n
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length. Apparently, there is no correlation of long Costa 8. Supra-antennal setae: (0) absent, (1) present ade
to R relative length, as there are taxa with long Costa d
with1 relatively short R –costal apex distance (e.g. (Fig. 10). C.I. = 10; R.I. = 73. fro
1 m
Dohrniphora Dahl) and with relatively long R –costal Phorids may have zero, one pair, or two pairs of supra- h
apex distance (e.g., Bothroprosopa Schmitz). H1owever, antennals. As all taxa with two pairs of supra-anten- ttp
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most taxa with short costa have R –costal apex dis- nals have these setae reclinated, I considered these ://a
1 c
tance shortened possibly due to dependence between conditions possibly related and coded the present a
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these conditions. This possible dependence was consid- character with only the distinction between supra- em
ered to apply this character only to taxa with costal antennals absent/present. The information on supra- ic.o
measuring 0.37 or more of the wing length. A short antennals inclination is dealt in character 9. up
R –costal apex distance was recovered as homologous Brown (1992) coded presence of supra-antennals .co
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in most taxa of the Thaumatoxena-group, with a rever- in character 2 of table 4 assigning the state of supra- /z
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sion optimized at the base of the Hypocera-subgroup. antennals present to all Euphorida. Brown et al. (2015) o
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considered this information only for hypocerines in n
e
character 76 ‘Supra-antennal setae of hypocerines: (0) an
OTher CharaCTers pthreemse nats, ‘(01’.) Haobwseenvte’,r ,t thhoeu vgahr isacboirliintyg otfa txhae epxrteesrennacel toof /article
7. Frontal furrow: (0) absent, (1) present (Fig. 9). these setae within Phoridae was not completely recov- /1
8
1
C.I. = 16; R.I. = 68. ered by the Bauplan approach of Brown (1992) and by /1
/1
5
1
/3
0
7
5
1
2
8
b
y
g
u
e
s
t o
n
1
0
A
u
g
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s
t 2
0
2
1
Figure 7. Examples of different conditions of character 5 (scutellum length/width).
Figure 8. Examples of different conditions of character 6 (distance R–costal apex/C length).
1
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
PHYLOGENY OF PHORINAE 159
D
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n
lo
a
d
e
d
fro
m
h
ttp
s
://a
c
a
d
e
m
Figure 9. Character 7 (frontal furrow). (A–D), examples of its states; (E), optimization in the strict consensus (colours rep- ic
resenting states: green 0; blue 1; pink missing, inapplicable, or ambiguous data). .ou
p
.c
o
Brown et al. (2015), as the absence of supra-antennals authors as a synapomorphy of Euphorida. However, m
/z
in taxa such as Aenigmatias and Termitophyllomyia genera that do not have ventral interfrontal setae o
o
Schmitz was not accounted in their coding. such as Chonocephalus, Cyphocephalus Borgmeier, lin
n
Supra-antennals were coded here for the entire fam- Thaumatoxena Breddin & Börner, Misotermes and ea
n
ily being optimized as a possible synapomorphy at the others were coded as having them. The pair of ventral /a
base of Metopininae + Phorinae, with nine reversions fronto-orbitals was coded in Brown’s (1992) analysis of rtic
le
occurring within these subfamilies, in some cases cor- ‘Hypocerinae’ as character 6 of table 5 ‘Lower fronto- /1
8
roborating large groups such as the reversion close to orbital seta absent’. Both of these characters were 1
/1
the base of the Hypocera-subgroup. united here in one with three conditions, which was /1
5
analysed for the entire family and treated as additive. 1
9. Supra-antennal setae inclination: (0) reclinate, (1) /3
The complete ventral row was recovered as a synapo- 0
proclinate (Fig. 11). C.I. = 100; R.I. = 100. 75
morphy of Phorinae + Metopininae. Nine reversions to 1
2
Supra-antennal setae reclinate has been recog- an incomplete ventral row corroborate small groupings. 8 b
nized as one of the clearest characters diagnosing 11. Inclination of frontal ventral row of setae: (0) y gu
the Metopinini (monophylum 4 in Brown, 1992). The e
approximately in the same horizontal line, (1) s
present character coded this information similarly angulated, with interfrontals shifted ventrally, (2) t on
to character 68 of Brown et al. (2015), except that 1
very angulated (Fig. 13). C.I. = 12; R.I. = 74. 0
here the character was applied only to taxa with the A
u
condition ‘Supra-antennals present’ (character 7). This character was applied only to taxa with all four g
u
Supra-antennals reclinate was corroborated as a syna- setae of the frontal ventral row (character 10, state st 2
pomorphy of Metopinini being recovered at the base of 2), and treated as additive (Fig. 13). Non-angulated or 0
2
1
Megaselia Rondani + Allochaeta Borgmeier. slightly angulated rows are optimized at the base of
large groups while very angulated row was optimized
10. Frontal ventral row of setae: (0) absent, (1) only
as a synapomorphy of groups of medium inclusiveness,
interfrontals present, (2) all four setae present
such as the grouping of Spiniphora Malloch species
(Fig. 12). C.I. = 18; R.I. = 68.
and of Phora + New amber genus P.
Phoridae ventral row of setae generally comprises a
12. Frontal dorsal row of setae: (0) absent, (1) only
pair of interfrontals (VIF) and a pair of fronto-orbit-
interfrontals present, (2) all four setae present
als (VFO). The pair of VIFs was coded in character
(Fig. 14). C.I. = 18; R.I. = 35.
64 of Brown et al. (2015) ‘Modern frontal setation
of phorids (ventral interfrontal setae present): (0) Phoridae dorsal row of setae generally comprises a pair
present, (1) absent’, which was recovered by these of interfrontals and a pair of fronto-orbitals. The three
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
160 D. C. AMENT
D
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a
d
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Figure 10. Character 8 (supra-antennal setae). (A–E), examples of its states; (F), optimization in the strict consensus (col- ://ac
a
ours representing states: green 0; blue 1; pink missing, inapplicable, or ambiguous data). d
e
m
ic
.o
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p
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o
m
/z
o
o
lin
n
e
a
n
/a
rtic
le
/1
8
1
/1
/1
5
1
/3
0
7
5
1
2
8
b
y
g
u
e
Figure 11. Character 9 (supra-antennal setae inclination). (A–C), examples of its states; (D), optimization in the strict s
consensus (colours representing states: green 0; blue 1; pink missing, inapplicable, or ambiguous data). t o
n
1
0
A
u
observed conditions of presence/absence of these pairs state 2). The state of interfrontals more dorsal than g
u
were treated here as an additive character (Fig. 14). fronto-orbitals was recovered as homologous in st 2
A complete frontal dorsal row of setae was optimized Diplonevra Lioy + Dohrniphora and possibly as a syn- 0
2
1
present in the most basal nodes of Phoridae, with inde- apomorphy of Hirotophora + Chaetocnemistoptera +
pendent losses occurring at the base of Chonocephalus Neopleurophora.
+ Termitoxeniinae, Dohrnigma Disney & Ellwood +
14. Additional frontal setulae: (0) absent, (1) present
Misotermes + Aenigmatistes and of some other taxa.
(Fig. S4, optimization). C.I. = 100; R.I. = 100.
13. Inclination of frontal dorsal row of setae: (0)
This character was used by Mostovski (1999) in the
approximately in the same horizontal line, (1)
investigation of the relationships among Sciadocerinae,
interfrontals more dorsal than fronto-orbitals
Prioriphorinae and Euphorida. In the present analy-
(Fig. 15). C.I. = 16; R.I. = 44.
sis, only Sciadocera and Archiphora do not have these
This character was applied only to taxa with all setulae, and their presence was recovered as homolo-
four setae of the frontal dorsal row (character 12, gous in all taxa that have this condition. However, this
© 2017 The Linnean Society of London, Zoological Journal of the Linnean Society, 2017, 181, 151–188
