A review of Cunaxidae (Acariformes, Trombidiformes): Histories and diagnoses of subfamilies and genera, keys to world species, and some new locality records Author Skvarla, Michael J. Department of Entomology, 319 AGRI Building, University of Arkansas, Fayetteville, Arkansas, 72701, USA Author Fisher, J. Ray Department of Entomology, 319 AGRI Building, University of Arkansas, Fayetteville, Arkansas, 72701, USA Author Dowling, Ashley P. G. Department of Entomology, 319 AGRI Building, University of Arkansas, Fayetteville, Arkansas, 72701, USA text ZooKeys 2014 2014-06-20 418 1 103 http://dx.doi.org/10.3897/zookeys.418.7629 journal article http://dx.doi.org/10.3897/zookeys.418.7629 1313-2970-418-1 D71C8A3DA6CA40A5B3A034A1FD1C16A0 F034FFA3FFAFFFBEB578C42DD65AFFA2 578347 Pulaeus Den Heyer, 1978 Historical review. Ewing (1909) described the first species of Pulaeus as Eupalus pectinatus . Berlese (1916) described Eupalus sternalis . Baker and Hoffmann (1948) proposed Cunaxoides to replace Eupalus as the name was preoccupied; described Cunaxoides patzcuarensis , Cunaxoides whartoni , and Cunaxoides americanus ; and synonymized Cunaxoides sternalis with Cunaxoides pectinatus . They also redescribed and illustrated Cunaxoides pectinatus . Muma (1960) described Cunaxoides pectinellus . Shiba (1978) described Cunaxoides neopectinatus , Cunaxoides parapatzuarensis , and Cunaxoides pseudominutus . Chaudhri, Akbar, and Rasool (1979) described Neocunaxoides krama . Kuznetzov and Livshitz (1979) reported Cunaxoides pectinatus and Cunaxoides americanus from Russia. Den Heyer (1979b) erected Pulaeus and moved the previously mentioned species into the new genus; he also redescribed Pulaeus pectinatus and described Pulaeus glebulentus . Neocunaxoides cinctus was described by Chaudhri (1980) . Den Heyer (1981c) confirmed the synonymy of Pulaeus sternalis with Pulaeus pectinatus , and synonymized Cunaxoides pectinellus with Pulaeus pectinatus ; he also described Pulaeus franciscae and placed Pulaeus within Cunaxoidinae , tribe Pulaeini . El-Bishlawy and Rakha (1983) described Pulaeus zaherii from Egypt. Liang (1983) reported Pulaeus pseudominutus from China. Pulaeus musci was described by Liang (1985) . Zaher and El-Bishlawy (1986) described Pulaeus niloticus . Bu and Li (1987b) described Pulaeus longignathos and Pulaeus chongqingensis . Muhammad and Chaudhri (1990) described Pulaeus desitis , Pulaeus ferventis , Pulaeus osculum , and Pulaeus verno from Pakistan. Pulaeus ardeola was described by Barilo (1991) . Muhammad and Chaudhri (1991a) described Pulaeus camar , Pulaeus erinaceus , Pulaeus galumma , Pulaeus haurio , Pulaeus silicula , and Pulaeus stultus from Pakistan. Smiley (1992) synonymized Pulaeus niloticus with Pulaeus subterraneus and provided a key to known world species; he also transferred Cunaxoides neopectinatus to Neocunaxoides . Li et al. (1992) recorded Pulaeus glebulentus from Chongqing, China. Corpuz-Raros (1996b) described two species, Pulaeus payatopalpus and Pulaeus rimandoi , from the Philippines. Lin and Zhang (2000) reported Neocunaxoides neopectinatus , Pulaeus longignathos , Pulaeus musci , and Pulaeus pseudominutus from China. Lin et al. (2003) reported Pulaeus minutus from China. Bashir, Afzal, and Akbar (2005) described Pulaeus punctatus . Bashir and Afzal (2006b) described Pulaeus anjumi . Corpuz-Raros (2007) also described Pulaeus cebuensis , Pulaeus palawanensis , and Pulaeus samarensis . Castro and Den Heyer (2009) split Lupaeus from Pulaeus and described two new species: Pulaeus myrtaceus and Pulaeus quadrisolenidius ; they also synonymized Pulaeus longignathos with Neocunaxoides krama and transferred Neocunaxoides krama to Pulaeus . Bashir and Afzal (2009) described Pulaeus akbari , Pulaeus banksi , and Pulaeus walii . Lin and Zhang (2010) argue that the "original species name longignathos [as in Pulaeus longignathos ] is the correct form in Greek. Some authors emended it to the Latinized form longignathus (e.g. Castro and Den Heyer, 2009: 2)." The spelling longignathos is followed here. Sergeyenko (2011b) described Pulaeus leonidi , Pulaeus maslovi , and Pulaeus semistriatus and synonymized Pulaeus longignathos and Pulaeus chongqingensis with Pulaeus krama as he considered them to be male and female of that species, respectively. Den Heyer et al. (2013) described Pulaeus razanensis . Diagnosis. Gnathosoma . Pedipalps 3-segmented. Femurogenua at least twice as long as wide, complemented with 6 setae. Tibiotarsi at least twice as long as wide, usually complemented with 6 setae, 1 pointed process, and may possess a bladder- or knob-like apophysis ( Fig. 39a-c ). Subcapitulum with 6 pairs of setae ( hg1 - 4 and 2 pairs of adoral setae); setae hg4 often the longest. Chelicera with seta present. Figures 35-39. Pulaeus illustrations. 35 Genital setae in a row 36-39 Pulaeus key illustrations 36, 37 Venter, setae removed for clairity 36a Coxae I-II not coalesced medially, median platelet present 36b Coxae I-II not coalesced medially, median platelet absent 37a Coxae I-II coalesced medially, median platelet present 37b Coxae I-II coalesced medially, median platelet absent 38a Dorsal shield with punctures 38b Dorsal shield with broken striae 39a-c Pedipalp tibiotarsus 39a Tibiotarsus with elongate apophysis 39b Tibiotarsus with flat apophysis 39c Tibiotarsus with flange-like apophysis. Idiosoma , dorsal . Proterosoma bears a well-sclerotized shield, complemented with 2 pairs of setae ( lps and mps ) and 2 pairs of setose sensillae ( at and pt ). Dorsal hysterosoma bears a sclerotized plate which is variable in size and fused with the proterosomal shield; it may be complemented with a variable number of setae depending on the size of the plate. Setae c1 - h1 , c2 , f2 , and h2 and present. Cupule im present laterad and posterior of e1 . Integument not covered in shields or plates striated. Idiosoma, ventral . Coxae sclerotized and well-defined. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV may be fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae ( g1 - 4 ), which are usually in a straight row; 2 pairs of genital papillae visible underneath the plates. Anal plates bear one pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. The integument not covered in shields or plates striated. Legs. Tarsi never constricted apically so as to end in lobes. Trichobothrium on leg tibia IV present. Depression of the famulus occurs on proximal half of tarsus I. Tibiae I-II possess non-striated blunt solenidia. Ambulacral claws rippled and occur on either side of a 4-rayed empodium. Key to adult female Pulaeus Pulaeus ardeola was not included in the key because the original text is in Cyrillic script and the illustrations do not provide enough characters to differentiate it from other species. Neocunaxoides cinctus is moved from Neocunaxoides to Pulaeus based on features given in the original description, namely that f2 is present and basifemora IV are complemented with 2 sts. The following were species assigned to Pulaeus before Lupaeus was erected. The characters that divide the two genera are not given in the original species descriptions and types have not been viewed. These indeterminable species are therefore not included in either generic key, but instead characters are given for each species that will serve to identify them. Pulaeus parapatzuarensis (Shiba, 1978) - This species has a divided sternal plate, lacks a sclerotized area anterior to the genital plates, and does not have f1,2 located on platelets. In addition it has 6 pairs of setae on the integument between coxal and genital plates. Pulaeus patzcuarensis (Baker & Hoffmann, 1948) - This species can be recognized by the sternal plates being connected anteriorly and divided in a v-shape posteriorly. Pulaeus pseudominutus (Shiba, 1978) - Setae e1 being 3 times the length of c1 and d1 distinguishes this species. Pulaeus payatopalpus (Corpuz-Raros, 1996) - The hypostome is 2/3 the length of the gnathosoma and the pedipalps are extremely long and slender, at least 8 times longer than wide. In addition the tibiotarsus is complemented with a seta that is longer than the segment. Pulaeus zaherii (El-Bishlawy & Rakha, 1983) - This species can be recognized by the divided sternal plates, f1 being 4/5 the length of e1 , and f1 being 1/2 the length of f2 .

1	Sternal plate divided medially ( Fig. 35a, b )	2
-	Sternal plate not divided medially ( Fig. 36a, b )	23
2 (1)	Median platelet between coxae II-III present ( Fig. 35a )	3
-	Median platelet between coxae II-III absent ( Fig. 35b )	7
3 (2)	Dorsal shield with surface smooth anteriorly and broken striae or lobes posteriorly; Ukraine	Pulaeus semistriatus Sergeyenko, 2011
-	Dorsal shield with surface patterned (broken striae/lobes or dotted) on entire surface	4
4 (3)	Dorsal shield patterend with dots; Pakistan	Pulaeus punctatus Bashir, Afzal & Akbar, 2005
-	Dorsal shield patterned with broken striae/lobes	5
5 (4)	Genua II with solenidia present	6
-	Genua II with solenidia absent; Pakistan	Pulaeus banksi Bashir & Afzal, 2009
6 (5)	Genua II with 1 asl, 5 sts; genua III with 2 asl, 5 sts; South Africa	Pulaeus glebulentus Den Heyer, 1979
-	Genua II with 2 asl, 4 sts; genua III with 1 asl, 5 sts; Iran	Pulaeus razanensis Den Heyer, 2013
7 (2)	Setae f1 and f2 located on sclerotized platelets or shields	8
-	Setae f1 and f2 not located on sclerotized platelets or shields	20
8 (7)	Pedipalp femurogenu at least 6 times as long as wide; Philippines	Pulaeus rimandoi Corpuz-Raros, 1996
-	Pedipalp femurogenu at most 4 times as long as wide	9
9 (8)	Genua II with 0 solenidia; Pakistan	10
-	Genua II with 1 solenidion	12
-	Genua II with 2 solenidia; Philippines	Pulaeus samarensis Corpuz-Raros, 2007
-	Genua II with 3 solenidia	17
-	Genua II with 4 solenidia	19
10 (9)	Genua I wth 2 bsl, 6 sts; tibia I with 1 bsl, 6 sts; Pakistan	Pulaeus ferventis Muhammad & Chaudhri, 1990
-	Genua I with 2 asl, 3 bsl, 3 sts; tibia I with 1 bsl, 7 sts; Pakistan	Pulaeus erinaceus Muhammad & Chaudhri, 1991
-	Genua I with 3 bsl, 6 sts; tibia I with 1 asl, 1 bsl, 6 sts; Pakistan	Pulaeus galumma Muhammad & Chaudhri, 1991
-	Genua I with 4 asl, 4 sts; tibia I with 1 asl, 6 sts; Pakistan	Pulaeus walii Bashir & Afzal, 2009
-	Genua I with 5 bsl, 4 sts; tibia I with 1 bsl, 6 sts; Pakistan	11
11 (10)	Basifemora I-IV setal formula 5-5-4-3; Pakistan	P. silicula Muhammad & Chaudhri, 1991
-	Basifemora I-IV setal formula 4-6-3-1; Pakistan	Pulaeus stultus Muhammad & Chaudhri, 1991
12 (9)	Basifemora I with solenidion present; telofemora I-IV setal formula 5-5-3-2; Pakistan	Pulaeus camar Muhammad & Chaudhri, 1991
-	Basifemora I with solenidion absent; telofemora I-IV setal formula not as above	13
13 (12)	Basifemora II with 5 (rarely 4) sts; Ukraine	Pulaeus leonidi Sergeyenko, 2011
-	Basifemora II with 6 sts	14
14 (13)	Genua II with solenidia present	15
-	Genua II with solenidia absent; Pakistan	Pulaeus akbari Bashir & Afzal, 2009
15 (14)	Genua II with 1 asl, 5 sts; Ukraine	Pulaeus maslovi Sergeyenko, 2011
-	Genua II with 1 bsl, 6 sts	16
16(15)	Genua III-IV setal formula 5 sts-5 sts; Pakistan	Pulaeus osculum Muhammad & Chaudhri, 1990
-	Genua III-IV setal formula 5 sts-6 sts; Pakistan	Pulaeus haurio Muhammad & Chaudhri, 1991
-	Genua III-IV setal formula 1 bsl, 4 sts-2 bsl, 4 sts; Pakistan	Pulaeus verno Muhammad & Chaudhri, 1990
17 (9)	Setae f1 and h1 approximately equal in length	18
-	Setae f1 approximately half the length as h1 ; China	Pulaeus musci Liang, 1985
18 (17)	Coxa IV with 2 sts; basifemora IV with 2 sts; Brazil	Pulaeus myrtaceus Castro & Den Heyer, 2009
-	Coxa IV with 3 sts; basifemora IV with 1 sts; Pakistan	Pulaeus anjumi Bashir & Afzal, 2006
19 (9)	Dorsal shield with punctuations ( Fig. 37a ); Brazil	Pulaeus quadrisolenidius Castro & Den Heyer, 2009
-	Dorsal shield with flat broken striae ( Fig. 37b ); USA	Pulaeus whartoni (Baker & Hoffmann, 1948
20 (7)	4 pairs of setae on integument between coxal and genital plates	Pulaeus cinctus (Chaudhri, 1980)
-	5 pairs of setae on integument between coxal and genital plates	21
-	6 pairs of setae on integument between coxal and genital plates	22
21 (20)	Coxae II with 2 sts; telofemora II with 5 sts; Pakistan	Pulaeus desitis Muhammad & Chaudhri, 1990
-	Coxae II with 2 sts; telofemora II with 4 sts; Philippines	Pulaeus palawanensis Corpuz-Raros, 2007
22 (20)	Sensillum at approximately as long as sce ; setae f1 approximately equal in length to h1	Pulaeus cebuensis Corpuz-Raros, 2007
-	Sensillum at longer than sce ; setae f1 approximately 1.25 the length of h1	Pulaeus franciscae Den Heyer, 1981
23 (1)	Ventral medial platelet present ( Fig. 36a ); dorsum punctuate ( Fig. 37a ); pedipalpal tibiotarsus with truncate, flange-like apophysis ( Fig. 38a ); USA	Pulaeus pectinatus Den Heyer, 1979
-	Ventral medial platelet absent ( Fig. 36b ); dorsum striated ( Fig. 37b ); pedipalpal tibiotarsus with elongate apophysis ( Fig. 38b )	24
24 (23)	Posterior pedipalpal tibiotarsal seta bifurcate ( Fig. 39 )	Pulaeus neopectinatus (Shiba, 1978)
-	Posterior pedipalpal tibiotarsal seta not bifurcate	25
25 (24)	Pedipalp femurogenua at most 4 times as long as wide; setae f1 and f2 approximately equal in length; USA	Pulaeus americanus (Baker & Hoffmann, 1948)
-	Pedipalp femurogenua at least 6 times as long as wide; setae f1 1/4 longer than f2 ; Pakistan	Pulaeus krama (Chaudhri, Akbar & Rasool 1979)