Diapherodes achalus (Rehn, 1904) Rehn, 1904

Frank H. Hennemann, Oskar V. Conle & Daniel E. Perez-Gelabert, 2016, Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “ Anareolatae ”: Phasmatidae: Cladomorphinae), Zootaxa 4128 (1), pp. 1-211 : 89-93

Diapherodes achalus (Rehn, 1904) n. comb.

( Figs. 157–166 , 350 , 370 , 382–383 )

Aplopus achalus Rehn, 1904: 68 . HT, ♂: Adjuntas P.R. 12 Apr. 0 0, C.W. Richmond Collector, Cat. no. Aplopus achalus Rehn

Type, Type No. 7344 U.S. N.M. [ USNM, No. 7344].

Wetmore, 1916: 58.

Wolcott, 1923: 23.

Wolcott, 1936: 35.

Wolcott, 1948: 50.

Wolcott, 1951: 50.

Diapherodes achalus, Van de Bussche et al., 1988: 422 . Haplopus achalus, Kirby, 1904a: 363 .

Otte & Brock, 2005: 150.

Diapherodes achalus, Moxey, 1972: 90 (in litt.). Diapherodes longiscapha Redtenbacher, 1908: 435 . LT (by present designation), ♀: Porto-Rico, Antilles, Mr. H. de Saussure,

Diapherodes longiscapha REDT. [ MHNG]; PLT, ♀: 141, Portorico Coll. Krug, Diapherodes longiscapha Br. Brunner det,

Type [MNHU] n. syn.

Wolcott, 1923: 23.

Wolcott, 1936: 35.

Wolcott, 1948: 50.

Wolcott, 1951: 50.

Zompro & Brock, 2003: 16.

Otte & Brock, 2005: 121.

Diapherodes gigantea, Wolcott, 1923: 23 . (Misidentification —not Mantis gigantea Gmélin, 1789 ) Wolcott, 1936: 35.

Wolcott, 1948: 50.

Wolcott, 1951: 50.

Diapherodes krugii, Wolcott, 1923: 23 . (Nomen nudum) Wolcott, 1936: 35.

Further material [16 ♂♂, 15 ♀♀, 1 nymph, 1 egg]:

PUERTO RICO:

1 ♂: Puerto Rico: Parador, Casa Grande, nr. Utuado, 16-III-1990, at light, R.E. & K.E. Wooruff [ FSCA]; 1 ♀: Barranguitas, P.R., June 1959, coll. L.F. Martorell [ USNM]; 1 ♀ (in Ryker mount): From Porto Rico Exp. Stat. […] Nov. 19.1913; Diapherodes longiscapha Redt. ♀ A.n.c. [ USNM]; 1 ♀: Road S.G. to Mariacao, 18K6M side road, Beating from Tree, Feb. 13,'61, A.M. Nadler; P.R. Road from Maricao-Sabana Grande km 18.6, Beating tree, Feb. 13,'61, A.M. Nadler; Diapherodes longiscapha Redt. ♀, det. A.B. Gurney 1961 [ USNM]; 1 ♂: P.R. ACC.NO. 69–50, Cayey, P.R., Sept. 26,1950; G.N. Wolcott & L.F. Martorell Collectors; USMNH; w0150; Diapherodes achalus (Rehn) det. CF Moxey 1972 [ USNM]; 1 ♂: Utuado, P.R., Nov. 1948, Coll: E. Boneta; w0152; USMNH; Diapherodes achalus (Rehn) det. CF Moxey 1972 [ USNM]; 1 ♂: Caguas, P.R., Jan. 8.1912, in bird stomach. Biol. Survey.; Aplopus achalus Rehn A. n.c [ USNM]; 2 ♂♂: Barranquitas, P.R., Nov. 20 1948, Coll. S. Rodriguez [ USNM]; 1 ♂: Mayaguez, P.R., 1/14/47, C. Feyre R. [ USNM]; 1 ♂: Mayaguez, P.R., 1-19-1930, Coll. E. Rosa [ USNM]; 1 ♂: Ponce, P.R., X-20 1947 [ USNM]; 1 ♀: Juyuya PR, VIII-14- 54, H F Winters, PR-1278; Toro Negro Ar, Carribean Nat. Forest, Lot 45-19676 [ ANSP]; 1 ♀ (valde defectum): Yauco, P.R., June 1945, Coll: [ USNM]; 2 ♀♀: Rio Piedras, P.R., IV-2-1944, Coll: R. Casas [ USNM]; 1 ♀: Barranquitas, P.R., Oct. 10 1948, Coll. S. Rodriguez [ USNM]; 1 ♀: Maricao, P.R., Nov. 20 1990, Coll: Iv. Nazario [ USNM]; 1 ♀: Aibonito, P.R., 8-9-33, R.G. Oakley; on leaf Inga vera ; San José No. 4630; wO143 [ ANSP]; 1 ♀: El Yunque Mountains, Luquillo, P.R., June 1935; cdl. L. F. Martore, Host: on weeds, Acc#: 29–35; wO144 [ ANSP]; 1 ♀: Jayuya, PR, Aug. 23–45, H F Winters, PR No. 472; on Cinchona ledgeriana ; Lot No. 56-1971; wO147 [ ANSP]; 1 ♂: Jayuya, P.R., Aug. 23–45, H F Winters, PR No. 470; on Cinchona ledgeriana ; wO151 [ ANSP]; 1 ♂: Mayaguez, PR; wO149 [ ANSP]; 1 ♀ (penultimate instar): Yauco, P.R., II.1934, Cdi J. Matini ; wO146 [ ANSP]; 1 ♀: Mayaguez, P. R,. 3. Oct. 1938; J. A. Ramos Collector [ ANSP]; 1 ♂: Mayaguez, P. R., 21 Oct. 1937, Coll.: R. del Moral [ ANSP]; 1 ♂: Aibonito, P. R., II-16-1935, Coll. J. Martinez [ ANSP]; 1 ♀, 1 egg (ex ovipositor): Porto Rico, Mayaguez, 5.7.1906, C. Gaggo leg.; PHA 12, Zoologisches Museum Hamburg [ ZMUH]; 1 ♀: Mayaguez, Porto Rico, C. Gaggo leg. 5.VII.1906, ded. 3.IX.1906; PHA 11, Zoologisches Museum Hamburg [ ZMUH]; 1 ♂: Puerto Rico, Maricao, 13.10.2009, leg. Mel J. Rivera [coll. OC].

NO DATA:

1 ♂: F. Sein Collector; Aplopus jamaicensis (Drury) A.n.c. [ USNM].

Diagnosis: Well characterized and distinguished from the two other species of the jamaicensis species-group by the elongate head, just gently rounded and almost unarmed vertex ( Figs. 159–160 ), entirely unarmed mesothorax ( Figs. 157–158 ) and having only two apical spines on the medioventral carina of the meso- and metafemora in both sexes. ♂♂ furthermore differ by the bold brown longitudinal median stripe on the tegmina ( Fig. 158 ) and brown cheeks ( Fig. 160 ). The characteristic eggs are unique within the entire genus by having a hollow polar area and a prominent conical swelling on the operculum ( Figs. 165–166 ).

Description: The colouration is described from several photos of live specimens taken at Maricao in Puerto Rico by Mel J. Rivera ( Figs. 374–375 ).

♀ ( Fig. 157 , 382 ). Fairly small (body length including subgenital plate 92.0–116.0 mm) and rather massive member of the jamaicensis species-group with a moderately widened mesothorax and abdominal segment VII. Colour bright apple green, the meso- and metapleurae ochre with a white longitudinal stripe along lower margin. Ventral body surface mostly whitish. Tegmina and costal region of alae bright green with the anterior margin darker in colour. Anal region of alae dull pink to violet. Antennae dark ochre dorsally and blackish brown ventrally. Eyes pale ochre.

Head: Elongate, ovate and 1.3x longer than wide, the vertex very gently convex and smooth except for a very faint pair of cephalad tubercles; the sinistral very faint ( Fig. 159 ). Eyes rather small, circular and their length contained a little more than 3x in that of cheeks. Antennae broken in all specimens at hand but at least reaching to metanotum. Scapus dorsoventrally compressed, constricted basally with the outer lateral margin gently rounded, 2.3x longer than wide. Pedicellus oval in cross-section about 2/3 the length of scapus.

Thorax: Pronotum considerably shorter and narrower than head, hardly longer than wide and gently narrowing towards the posterior. Transverse median depression deep but very short and curved. Surface unarmed except for a pair of small tubercles just in front and behind the median sulcus ( Fig. 159 ). Mesothorax about 1.6x longer than head and pronotum combined, constricted in anterior portion and then gradually widened towards the posterior; posterior margin more than 3x wider than anterior margin. Mesonotum very slightly tectiform longitudinally and widened towards the middle, then roughly parallel-sided; surface smooth except for a few scattered small tubercles, in particular in the anterior portion. Mesopleurae with a marginal row of spiniform tubercles or short spines; metapleurae armed with a row of about eight short spines. Pro-, meso- and metasternum smooth; the mesosternum occasionally with a small pair of granules posteriorly. Tegmina oval, reaching to posterior margin of metanotum and with the central protuberance moderate. Alae roughly equal in length to tegmina and reaching about half way along median segment.

Abdomen: Median segment slightly longer than wide and very gently widening towards the posterior; smooth. Abdomen strongly swollen sub-basally with segment II widening, IV narrowing and III widest. II–IV transverse and III almost 1.5 wider than long. V–VIII longer than wide, VII moderately expanded and with the lateral margins rounded ( Fig. 162 ). Tergites II–IV smooth, V–VII with four roughly parallel, longitudinal carinae which are most decided on VII ( Fig. 162 ). Sternites II–VII smooth except for a fine carina along lateral margins. Praeopercular organ formed by a brown tubercle close to posterior margin of sternum VII ( Fig. 350 ). Tergites VIII–X narrower than previous segments and roughly of uniform width. VIII about ¾ the length of VII and very slightly constricted medially, about 1.2x longer than wide; IX indistinctly longer than wide. Anal segment about equal in length to IX, with a slight longitudinal median carina in basal portion and flattened towards the posterior; posterior margin rounded and with a very small and shallow median indentation ( Fig. 162 ). Epiproct small and semi-circular with a keel dorsally ( Fig. 162 ). Cerci very small, round in cross-section and tapered towards a pointed apex. Subgenital plate elongate, lanceolate, longitudinally keeled and variable in length; projecting over apex of abdomen by more than the combined length of tergites IX–X ( Figs. 161–162 ).

Legs: Profemora slightly shorter, mesofemora about ¾ the length of mesothorax and metafemora reaching about half way along abdominal segment IV. Medioventral carina of all femora with two rather small spines in apical portion. Posteroventral carina of meso- and metafemora with one, the anteroventral carina with two distinct, black-tipped sub-apical spines. Basitarsi no longer than following two tarsomeres combined.

♂ ( Figs. 158 , 383 ). Small (body length 60.0–74.0 mm) and fairly robust for the jamaicensis -group with a very characteristic colouration, body surface glabrous, with very long alae (36.8–42.0 mm). General colour bright green, washed with brown, the lateral surfaces of the head, entire pronotum, mesonotum, mesopleurae, posterior portion of metapleurae and bases of all femora ochraceous. Abdominal tergites V–X green laterally, yellow dorsally and V–IX with a mid to dark brown longitudinal median stripe ( Fig. 321 ). Ventral surface of abdomen with a whitish wash. Head bright green with the cheeks ochraceous brown and two bold, washed yellow markings between the eyes ( Fig. 160 ). Anterior margin of tegmina and basal portion of the same region of the alae white, tegmina with a further broad brown longitudinal stripe, the innermost portion green with darker green veins and basal portion yellow ( Fig. 158 ); costal region of alae plain green with dark green longitudinal veins, the anal region very pale and translucent pink. Eyes mid to dark brown. Antennae greyish ochre. Ventral sub-apical spines of the meso- and metafemora black.

Head: Ovate and only about 1.2x wider than long, otherwise generally as in ♀♀; cephalad tubercles very faint ( Fig. 160 ). Eyes of moderate size and projecting hemispherically, their length contained less than 2x in that of cheeks. Antennae projecting over posterior margin of abdominal segment II and moderately robust; with 36–37 antennomeres. Scapus oval in cross-section, roundly rectangular with the inner margin concave and about 1.3x longer than wide. Pedicellus cylindrical and about 2/3 the length of scapus.

Thorax: Pronotum slightly shorter and narrower than the head, with the anterior portion slightly broader than posterior portion; median transverse depression distinct, gently curved and almost reaching lateral margin of segment. Surface smooth or with a very few minute granules ( Fig. 160 ). Mesothorax 1.8x longer than head and pronotum combined; mesonotum at best with a few very obsolete granules anteriorly. Mesopleurae smooth, metapleurae with a few small granules. Meso- and metasternum smooth. Tegmina oval and with a rather longitudinal and acute central protuberance, roughly reaching to posterior margin of metanotum. Alae ± reaching to posterior margin of abdominal tergum VII ( Fig. 158 ).

Abdomen: Median segment gently narrowed towards the posterior. Segments II–VI slightly decreasing in length and width, II 2.7x and VI only about 2x longer than wide. elongate, cylindrical, parallel-sided and gradually decreasing in length. II 3.5x, VII 2.5x longer than wide. VII about equal in length to VI and very slightly widened posteriorly. Sternites II–VII smooth. Tergum VIII about 3/5 the length of VII, widened towards the posterior and slightly tectate. IX slightly shorter than VIII and very slightly narrowed towards the posterior. Anal segment about as long as IX with two longitudinal carinae dorsally, slightly constricted towards the posterior and the posterior margin sub-truncate with a shallow median emargination ( Fig. 164 ). Vomer roughly triangular with a moderately long terminal hook ( Fig. 370 ). Cerci about 2/3 the length of anal segment, rather slender, cylindrical and very slightly in-curving with the apex blunt. Poculum moderately convex, cymbiform and longitudinally carinate, the posterior margin rounded and with a narrow median emargination; very slightly projecting over posterior margin of tergum IX ( Fig. 163 ).

Legs: Shape and armature generally as in ♀♀.

Egg ( Figs. 165–166 ): Medium sized (capsule length 5.0 mm), capsule about 2x longer than wide and slightly higher than wide. Capsule surface srtrongly and irregularly sculptured with several raised swelling and an irregular swollen keel indicated, that surrounds the micropylar plate and posteriorly runs towards lateral surfaces of capsule. Another distinctly raised, blunt keel below micropylar plate, which almost reaches to the polar area. The polar area hollow, impressed and surrounded by an irregularly toothed rim. Anterior margin with an irregularly crown-like rim of tooth like and anteriorly directed swellings. Micropylar plate small, heart-shaped and its length contained about 3.5x in that of capsule. Central portion impressed but otherwise strongly swollen. Operculum sub-circular with a large, irregularly sculptured conical swelling in centre. General colour pale to mid ochraceous to reddish brown.

Measurements [mm]: Length 5.0, length (including operculum) 6.2 width 2.5, heigth 2.9, length of micropylar plate 1.3.

Comments: Moxey (1972: 10, in litt.) stated D. achalus to have been taken on Guarea ramiflora (Meliaceae) and Inga vera (Mimosaceae) which might both be part of its natural diet. Body lengths cited by Moxey (1972: 92, in litt.) are 61.0–74.0 mm for ♂♂ and 92.0–114.0 mm including the subgenital plate for ♀♀.

Distribution: Puerto Rico: Barranquitas [USNM]; Mayaquez [ANSP, ZMUH]; Maricao Forest [ANSP, coll. OC]; Yauco [ANSP]; between Maricao and Sabana Grande km 18.6 [USNM]; Cayey [USNM]; Adjuntas [ANSP, USNM]; Utuado [USNM]; Jayuya [ANSP]; Aibonito [ANSP]; Ponce [USNM]; Rio Piedras [USNM]; Caguas [USNM]; Naguabo, El Yunque Quadrangle, Naguabo [Van den Bussche et al., 1988: 423] & Luquillo Experimental Forest, El Verde [ANSP]. Endemic.

Number of specimens examined: 35

TABL 16. Measurements of Diapherodes achalus (Rehn, 1904) n. comb. [mm] ......continued on the next page TABL 16. (Continued)

Figures

FIGURES 157 – 160. Diapherodes achalus (Rehn, 1904). 157. ♀: Puerto Rico, Mayaguez [ANSP]; 158. ♂: Puerto Rico, Parador Casa Grande [FSCA]; 159. ♀ head and pronotum: Puerto Rico, Mayaguez [ANSP]; 160. ♂ head and pronotum Puerto Rico, Parador Casa Grande [FSCA].

FIGURES 161 – 166. Diapherodes achalus (Rehn, 1904). 161. ♀ apex of abdomen of synonym HT of D. longiscapha (lateral view) [MHNG]; 162. ♀ apex of abdomen of HT of synonym D. longiscapha (dorsal view) [MHNG]; 163. ♂ apex of ♂ abdomen (lateral view) [FSCA]; 164. ♂ apex of abdomen (dorsal view) [FSCA]; 165. Egg: Puerto Rico, Mayaguez (dorsal view) [ZMUH]; 166. Egg: Puerto Rico, Mayaguez (lateral view) [ZMUH].

FIGURES 349 – 356. Ventral view of abdominal sternum VII of ♀♀ showing the praeopercular organ and anal segment of ♂♂ showing the vomer. 349. Diapherodes martinicensis Lelong & Langlois, 2005 ♀: captive reared from Martinique [coll. FH, No. 0750 - 1]; 350. Diapherodes achalus (Rehn, 1904) ♀: Puerto Rico, Mayaguez [ANSP]; 351. Diapherodes jamaicensis (Drury, 1773) ♀: captive reared from Jamaica [coll. FH, No. 0428 - 17]; 352. Aploploides stenocephalum Rehn & Rehn, 1938 ♂ PT: NW-Cuba, Pinar del Rio [ANSP]; 353. Cephaloplopus euchlorus n. gen., n. sp., ♂ PT: Dominican Republic, Santo Domingo Province, Sierra Prieta [USNM]; 354. Cephaloplopus pulchellus n. gen., n. sp., ♂ HT: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 355. Parhaplopus evadne (Westwood, 1859) n. gen., n. comb. ♂ HT: Dominican Republic, Santo Domingo [NHMUK]; 356. Parhaplopus navarroi n. gen., n. sp. ♂ HT: Dominican Republic, Monseñor Nouel Province, Bonao [USNM].

FIGURES 365 – 372. Ventral view of anal segment of ♂♂ showing the vomer. 365. Diapherodes angulata (Fabricius, 1793) ♂: Guadeloupe [MNHN]; 366. Diapherodes dominicae Rehn & Hebard, 1938 V: Guadeloupe, Picard River track [coll. ASPER]; 367. Diapherodes gigantea saintluciae n. ssp. ♂ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 368. Diapherodes gigantea gigantea (Gmélin, 1789) ♂: captive reared from Grenada [coll. FH, No. 0359 - 34]; 369. Diapherodes martinicensis Lelong & Langlois, 2005 ♂: captive reared from Martinique [coll. FH, No. 0750 - 5]; 370. Diapherodes achalus (Rehn, 1904) ♂: Puerto Rico, Parador, Casa Grande [FSCA]; 371. Diapherodes jamaicensis (Drury, 1773) ♂: Jamaica, Portland [FSCA]; 372. Diapherodes laevicollis Redtenbacher, 1908, ♂: Jamaica, Saint Andrew [FSCA].

FIGURES 382 – 384. Live insects. 382. Diapherodes achalus (Rehn, 1904) ♀: Puerto Rico, Maricao [Ó Mel J. Rívera]; 383. Diapherodes achalus (Rehn, 1904) ♂: Puerto Rico, Maricao [Ó Mel J. Rivera]; 384. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀: captive reared from NW-Cuba, Pinar del Rio.

FIGURES 373 – 374. Maps. 373. Map of the West Indies showing the principle islands; 374. Known distribution of Aploploides Rehn & Rehn, 1938 [green] and Venupherodes n. gen. [red].

FIGURES 375 – 376. Maps. 375. Known distribution of Diapherodes Gray, 1835 and the two species-groups here defined: Gigantea - group [green] and Jamaicensis - group [red]; 376. Distribution of Haplopus Gray, 1835. Although there have not yet been definite records from the southern islands of the Bahamas Caicos and Turks Islands, Haplopus is most certainly also present on these islands. Hence, they are here included in the distribution range.

FIGURES 315 – 321. Parhaplopus navarroi n. gen., n. sp .. 315. Apex of abdomen of ♀ PT (lateral view) [USNM]; 316. Apex of abdomen of ♀ PT (dorsal view) [USNM]; 317. Apex of abdomen of ♂ HT (lateral view) [USNM]; 318. Apex of abdomen of ♂ HT (dorsal view) [USNM]; 319. Right tegmen and ala of ♂ HT [USNM]; 320. Egg PT (dorsal view) [USNM]; 321. Egg PT (lateral view) [USNM].