CHINESE PHLEBOTOMINE SANDFLIES
OF S U B G E N U S ADLERIUS N I T Z U L E S C U , 1931 ( D I P T E R A : PSYCHODIDAE) A N D THE IDENTITY OF PHLEBOTOMUS SICHUANENSIS LENG & YlN, 1983
Part I - Taxonomical study and geographical distribution
L E N G Y A N J I A * & Z H A N G L I N G M I N *
S u m m a r y :
Four species of Adlerius phlebotomine sandflies have been recorded in China, namely: P. chinensis Newstead, 1 9 1 6 (Pc); P.
fengi Leng & Z h a n g , 1 9 9 4 ; P. longiductus Parrot, 1 9 2 8 and P.
sichuanensis Leng & Y i n , 1 9 8 3 (Ps), Adlerius phlebotomines are the main vectors of visceral leishmaniasis (VL) in China; three of them are acknowledged as VL vectors and P. fengi is considered a potential VL vector for southwestern mountainous region.
Different opinion has been raised to the validity of identity of Ps by some investigators from Shanghai and Shanxi who consider Ps to be a large type of Pc instead of an isolate species. The center of controversy is whether Ps is an isolate taxon or a large type of Pc. The present authors have carried out a series of comparative studies for these two flies on: 1 quantitative and qualitative morphological characters of four Chinese Adlerius phlebotomines;
and 2 . differences in geographical distribution. All specimens of Pc and Ps used in the present study are collected where their holotypes-paratypes were produced - W e s t Mountain, W e s t Suburb, Beijing and Lixian County, Sichuan Province.
The results have forcefully proved that Ps is an isolate species instead of a so-called large type Pc according to the concept of species. The clarification of their taxonomical identities is meaningful because both of them are VL vectors in different epidemic areas in China; especially Ps is an important VL vector in high mountainous regions of southwestern China and some extend to the Loess Plateau of northwestern China, where VL still exists and it is also the first Phlebotomine sandfly discovered in Tibet, the locality being near Assam in India (Leng et al. 1 9 9 0 ) .
KEY WORDS : taxonomy, Diptera, Phlebotomus, Adlerius, P. sichuanensis.
MOTS CLES : taxonomie, Diptera, Phlebotomus, Alderius, P. sichuanensis.
RESUME : LES PLÉBOTOMES DE CHINE DU SOUS-GENRE ADLERIUS NITZULESCU, 1931 (DIPTERA : PSYCHODIDAE) ET IDENTITÉ DE PHLEBOTOMUS SICHUANENSIS LENG & YIN, 1983. I : TAXONOMIE ET DISTRIBUTION GÉOGRAPHIQUE
Quatre espèces de phlébotomes du sousgenre Adlerius sont connues de Chine. Elles sont respectivement nommées : P.
chinensis Newstead, 1916 (Pc), P. fengi Leng & Zhang, 1994; P.
longiductus Parrot, 1928 et P. sichuanensis Leng & Yin, 1983 (Ps). Trois d'entre elles sont incriminées dans la transmission de la leishmaniose viscérale en Chine et P fengi est considéré comme vecteur potentiel dans le sud-ouest montagneux.
La validité du taxon P. sichuanensis a été remise en question par les chercheurs de Shangai et Shanxi (ISS). Ils considèrent Ps comme un variant de grande taille de P. chinensis plutôt qu'une espèce à part entière. Cette différence d'identification est devenue le centre d'une controverse. Le présent travail est basé sur une série d'études comparatives de ces deux insectes.
Les études morphologiques, onatomiques et morphométriques des quatre espèces du sous-genre Adlerius de Chine sont rapportées ainsi que les différences de distribution géographique. Tous les échantillons de Pc et Ps utilisés dans cette élude ont été collectés dans les localités des holotypes et paratypes, région montagneuse de l'ouest, région sud-ouest (Shangai), région de Pékin et Lixian et la province de Sichuan.
Les résultats montrent clairement que Ps est une espèce en soi et non un représentant de Pc de grande taille. La clarification de la position taxonomique des deux espèces est fondamentale en raison de leur responsabilité dans la transmission de la leishmaniose viscérale épidémique dans les différentes localités chinoises.
Ps est un vecteur important de VL dans les régions de haute montagne du sud-ouest de la Chine et quelques foyers du plateau de Loess du nord-ouest où la VL existe encore. Par ailleurs c'est dans cette région que le premier phlébotome tibétain a été découvert localisé près d'Assam en Inde (Leng et al., 1990).
ABBREVIATIONS:
Pc: Phlebotomus chinensis, Ps: Phlebotomus sichuanensis, PS: phlebotomine sandfly, VL: visceral leishmaniasis, ISS: investigators from Shanghai Institute of Parasitic Diseases, Chi- nese Academy of Preventive Medicine, Shanghai and Shanxi Medical University, Taiyuan, Shanxi Province.
INTRODUCTION
* Department of Medical Parasitology, Faculty of Medicine, Jinan Uni- versity Guangzhou, 510632 China.
Correspondence: Leng Y.J.
Tel.: +86-20-85220255 or 85220270 - Fax: +86-20-85221343.
E-mail: 85220255@163.net
L
eng & Yin ( 1 9 8 3 ) reported 10 s p e c i e s o f PS from northwestern Sichuan, including two n e w species among which Phlebotomus sichuanensis (Ps) was described for the first time. Since then Ps has b e e n incriminated as a vector o f VL in Shanxi, Gansu, Qin- ghai and Sichuan Provinces ( Y i n & Leng, 1 9 8 5 ) . This sandfly is also distributed in Y u n n a n Province and Tibet A u t o n o m o u s Region. (Leng, Zhang & Zhang, 1 9 9 1 ; Leng, 1 9 9 7 ) . Xiong, Jin, Hong et al. ( 1 9 9 0 ) syno- nymized Ps to Phlebotomus chinensis (Pc) and consi-Parasite, 2001, 8, 3-9 3
Article available athttp://www.parasite-journal.orgorhttp://dx.doi.org/10.1051/parasite/2001081003
LENG YAN JIA & ZHANG LING MIN
dered Ps to b e a large type o f Pc. But they have not given any description to define a measurable distinc
tion b e t w e e n so-called small a n d large type Pc e x c e p t they stressed o n the altitude w h e r e the s p e c i m e n s are collected, at a b o v e 2 , 0 0 0 m or b e l o w 1,500 m o f s e a level b e c a u s e they c o n s i d e r e d that increasing altitude m a d e usual Pc c h a n g i n g into a large type Pc and this changing might reverse (Xiong, J i n , H o n g et al, 1 9 9 0 ) . T o g o further, ISS carried out a series o f study c o n c e r ning this item, they confirmed that Ps is a large type o f Pc instead o f an isolate taxon b a s e d on the follo
wing reasons: 1. large and small type i m a g o e s o f Pc w e r e c o l l e c t e d from different heights ( 9 8 0 - 2 , 3 7 0 m alti
tude) a n d t h e fourth instar h a t c h e d from them in labo
ratory are identical in m o r p h o l o g y a n d measurement ratios. 2. T h e male ascoid formula o f Pc has t w o pat
terns, 2 / 3 - 1 5 a n d 2 / 3 - 8 , 1/9-15; following t h e increa
sing altitude "the higher the altitude the m o r e the latter pattern ratio", u p to 9 6 . 6 % at a b o v e 2 , 0 0 0 m sea level.
Large type is often distributed at a b o v e 1,500 m s e a level ( X i o n g & J i n et al, 1 9 9 0 ) . 3. Zhao, Li et al. ( 1 9 8 7 ) and Zhao, Q i a o et al ( 1 9 9 8 ) c o m p a r e d the SEM ima
gines o f female pharyngeal armature - a subgenus cha
racter for Acileriiis instead o f inter-species characters - o f large a n d small type Pc a n d according to its simi
larity c o n c l u d e d that Ps is a large type Pc.
MATERIALS AND METHODS
O
rigin o f s p e c i m e n s and materials: for the present research Ps s p e c i m e n s w e r e c o l l e c t e d b y aspirator from a b o v e 9 0 0 m altitude, Lixian County, northwestern Sichuan, w h e r e its holotype- paratypes w e r e c o l l e c t e d for the first time. Pc speci
m e n s w e r e collected from: 1. T e m p l e s in West Moun
tain, West Suburb, Beijing w h e r e its h o l o t y p e w a s c o l l e c t e d in 1 9 1 5 ; 2. Lixian C o u n t y a n d N a n p i n g County, northwestern Sichuan, in the latter w h e r e both Ps and Pc are present. P.fengi s p e c i m e n s m e a s u r e d are their holotype and paratypes preserved in J i n a n University (Leng & Zhang, 1 9 9 4 ) a n d the measurement materials o f P. longiductus are q u o t e d from original descriptions a n d well k n o w n papers (Parrot, 1 9 2 8 ;
W a n g et al., 1966; Artemiev, 1 9 8 0 ; Lewis, 1982; Feng, 1 9 8 3 ) . T h e t a x o n o m i c a l systems o f Artemiev ( 1 9 8 0 ) , Lewis ( 1 9 7 7 , 1 9 8 2 , 1 9 8 7 ) a n d T h e o d o r ( 1 9 5 8 ) are adopted.
Quantitative and qualitative morphological study: c o m parative morphological structures are studied under m i c r o s c o p e . Its c o r r e s p o n d i n g imagines are drawn under c a m e r a lucida and m e a s u r e d in um scale for quantitative analysis.
Zoogeographical distribution: according to the data stu
died b y the present author (Leng, 1 9 9 7 ) .
RESULTS
MORPHOLOGICAL CHARACTERS
B
y comparing o f ten morphometric a n d t w o mor- phostructural characters o f males a n d four morphometric a n d t w o morphostructural c h a r a c ters o f females for t h e four Adlerius PS in China ( T a b l e s I, II, III a n d Fig. 2 for c o m p a r i s o n b e t w e e n Ps and Pc), the following facts are found:
Morphometrically: 1. Ps is o n e o f the largest PS in China a n d over the world. 2. A3, A3/L, ascoid o n A4, style a n d a e d e a g u s tubercle distance o f Ps is the lar
gest but its c / b is the smallest a m o n g the s p e c i e s in s u b g e n u s Adlerius ( T a b l e s I, II). 3. A4 in Ps is m u c h longer than that o f Pc [male = 7.6:1 ( 2 5 7 : 3 4 ) , female = 2.9:1 (148:51)]. 4. T h e male wing length is much longer in Ps [Ps/Pc = 1.5:1 ( 3 , 1 8 0 : 2 , 1 0 4 ) , T a b l e III]. 5. A3/L:
c5Ps > o*Pc, $Ps < 9 P c (Tables I, II). 6. Aedeagus and genital filament: Ps>Pc. 7. T h e a e d e a g u s tubercle dis
tance o f Ps is the longest o f the four (Ps: Pc = 1.4;
Ps/P. longiductus = 2.6; P s / P . f e n g i = 4 . 2 ) . 8. Genital filament/Genital p u m p = Ps>Pc ( 7 . 1 : 5 . 9 ) . 9. T h e fol
lowing characters o f Ps are o p p o s i t e to the other three Chinese Adlerius p h l e b o t o m i n e s : 1. Male A4 in Ps a n d P.fengi are m u c h longer (6/2 = 1:0.58 a n d 1:0.88, T a b l e III) instead o f shorter than that o f females;
2. Wing o f male Ps is longer instead o f shorter than that o f females ( 6 7 $ = 1:058, T a b l e III).
Morphostructurally: Ps differs from Pc in 1. Ps has n o c o m m o n spermathecal duct. 2. Male ascoid formula
5 A3 A3/L AF HC FHC ST GF Style Aed F / P
P c 419 1.59 2/3-15 24 0.66 31 679 202 175 5.9
I'f 460 1.47 2/3-8,15 1/9-57 0.59 II) 1,333 206 233 10.5
PI 417 1.52 2/3-8,15 1/9-64 0.57 16 1,400 2<>i 195 10.1
Ps 550 1.67 2/3-8,15 1/9-39 0.71 42 1,130 250 190 7.1
Pc: P. chinensis; Pf: P.fengi; PI: P. longiductus; Ps: P. sichuanensis.
A3: antenna 3; A3/L: antenna 3/labrum; AF: ascoid formula; HC: hairs on inner surface o f coxite; FHBC: farmost hair from base of coxite;
ST: subapical tubercle of aedeagus; GF: genital filament: Aed: aedeagus; F/P: genital filament/genital pump.
All measures in um.
Table I. - Some morphological characters of four Chinese Adlerius phlebotomines ( o* ).
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CHINESE PHI.EBOTOMINE SANDFLIES
9 A3 A 3 / L CDS SS
P. cbinensis 325 1.27 + 1 3 - 1 4
P. fengi 455 1.38 -
P. longiductus 388 1.06 + 11-13
P. sichuanensis 370 1.02 15-18
CDS: common duct of spermatheca; SS: segments of spermatheca.
Tabic* II. - Some morphological characters of four Chinese Adlerius phlebotomines (9)
Antenna 4 Wing length
Species S 9 a/9 S 9 ( J / 9
P. cbinensis 34 51 1:1.50 2.104 2.320 1:1.10 P. fongi 203 178 1:0.88 2,587 3.116 1:1.20 P. longiductus 56 66 1:1.18 2,363 2.707 1:1.15 P. sichuanensis 257 148 1:0.58 3,180 2.750 1:0.86
Table III. - Length of antenna 4 and wings of four Chinese Adle
rius.
Fig.I. - Map of the zoogeographical distribution of four Chinese Adlerius phlebotomine sandflies.
(Table III) in Ps is 2 / 3 - 8 , 1/9-15 which differs from that o f Pc 2 / 3 - 1 5 . Hundreds o f Ps male specimens collected from Y u n n a n , Tibet and Sichuan are repeatedly e x a mined: it s h o w s that all male Ps have only o n e pat
tern o f ascoid formula without variations. 3- Hairs o n the inner surface o f c o x i t e in Ps c o v e r e d a wider area and are m o r e in n u m b e r (Ps:Pc = 3 9 : 2 4 ) . T h e a b o v e m o r p h o l o g i c a l characters m a k e up sharp s p e c i e s dis
tinction b e t w e e n Ps and Pc ( T a b l e s 1. II, III).
ZOOGEOGRAPHICAL DISTRIBUTION
T h e t o p o g r a p h y o f China by according to height is clearly divided into three platforms o f steps. T h e first platform c o v e r s a h u g e area from the coast o f Pacific O c e a n w h e r e it is l o w e r than 5 0 0 m altitude. T h e s e c o n d platform c o v e r s w e s t e r n loess plateau and w e s t e r n m o u n t a i n o u s area w h e r e it is l o w e r than 1,000 m altitude. T h e third platform c o v e r s southwes
tern mountainous region higher than 1,000 m altitude including Tibet, the r o o f o f the Earth. P. cbinensis is mainly distributed in the first platform up to certain depth into the b o r d e r o f s e c o n d platform. P. sichua
nensis is distributed in s e c o n d and third platforms from
9 0 0 m to 2,800 m altitude with s o m e overlaps with P. cbinensis in the border o f s e c o n d platform. This has b e e n well studied for years (Leng, 1997; Zhang & Leng, 1997; Fig. 1 ) .
DISCUSSION
SPECIMENS USED IN THE STUDY
I
n the studies o f ISS, all p h l e b o t o m i n e s used w e r e said to be: "The locality o f specimens collected" that is in a limited area in northwestern Sichuan, southern Shaanxi and Shanxi (Fig. 1), just at/near to the places in a parapatric z o n e for Pc and Ps. But, from the contradictory descriptions given by them (Table IV), people can easily find that the origin o f specimens used in their works are mixed and blurred. ISS have never collected s p e c i m e n s from the p l a c e w h e r e original holotype or paratype o f Pc and Ps were formally col
lected.
T h e standard considered by ISS to differentiate Ps and Pc are: 1. T h e Size o n w h i c h d e p e n d s identification
S a m e p h o t o s Zhao, Li, Qiao & Y | n ( 1 9 9 7 ) # Zhao, Qiao. Y i n . Yin, Li ( 1 9 9 8 ) # 1 (1997) = D (1998)
2 (1997) - A (1998) 3 (1997- - C (1998) 4 (1997) = E (1998)
Large type P c from Nanping, Sichuan Province (2,400 x ) Small type P c from Nanping, Sichuan Province (1,600 x ) P c from Jixian. Shaanxi Province (2,000 x)
P c from Yichuan, Shaanxi Province (1,300 x )
Pc from Yichuan 2, Shaanxi Province
Large type P c from Nanping, Sichuan Province P c from Yichuan, Shaanxi Province
P c from Jixian. Shanxi Province
** The names underlined are the same writer's in both papers.
Table IV. - The same SEM imagine illustrated in different origin of places and species.
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L E N G Y A N J I A & Z H A N G L I N G M I N
Fig. 2. - P. chinensis. 1. genitalia; 2. aedeagus lateral view; 3, spermatheca; 4, pharynx (after Theodor, 1958); P. sichanensis. 5, genitalia;
6, aedeagus, lateral wiew and ventral view; 7, paramere showing the short hairs in the middle of lateral side; 8, spermatheca; 9, furca;
10, pharynx viewed at three different focal depths for the same specimen (after Leng & Yin, 1983).
6 Parasite, 2001, 8, 3-9
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CHINESE PHLEBOTOMINE SANDFLIES
o f large and small type Pc b y these investigators is pro
bably corresponding to a microscopical aspect without distinctive data for size distinction. 2. T h e height is c o n s i d e r e d to b e an important factor for dividing Ps and Pc b e c a u s e Investigators o f Shangai Institut consi
der, that following the increasing altitude, the size o f p h l e b o t o m i n e sandflies is enlarging. W h e n r e a c h e d 1,500-2,000 m in altitude large type Pc appeared. S o the s p e c i m e n c o l l e c t e d a b o v e or lower 1,500 m w e r e treated as large a n d small Pc respectively for their stu
dies. But w e have found Ps a p p e a r e d in the height o f 900 m a b o v e sea level in northwestern Sichuan, 2,800 m in Y u n n a n a n d 2 , 5 0 0 m in Tibet.
Most proper morphological differences useful for iden
tifying these t w o flies described in the world literature under the light o f contemporary taxonomy do not s e e m to have b e e n c o n s i d e r e d b y ISS.
T h e s p e c i m e n s used for their studies c a m e from a similar origin ( X i o n g & Lu's old dry preserved speci
m e n s ) . Upon the facts m e n t i o n e d a b o v e it is hard to avoid w r o n g identification a n d confusions for the ori
ginal s p e c i m e n s used in their work.
X i o n g , J i n , Hong et al. ( 1 9 9 0 ) stated in their p a p e r that
"We c h e c k e d the s p e c i m e n s sent to us as a gift b y D r Yin Zhi-cheng o f Sichuan Institute in Parasitic Diseases c o l l e c t e d from Huan (Lixian?) County at 2,400 m and 2,800 m altitude. T h e n these very large specimens w e r e r e n a m e d b y Leng et al. (It is really Leng Y . J . & Y i n Z.C., 1 9 8 3 ) as P. sicbuanensis, w h i c h is actually what w e h a v e repeatedly e x p l a i n e d the large type P. chi-
nensis distributed 2 , 0 0 0 m a b o v e sea level. S o that P. sicbuanensis should b e a s y n o n y m o f P. cbinensis."
Dr Y i n Zhi-cheng c o l l e c t e d a large n u m b e r o f PS in an area w h i c h c o v e r s 2 9 - 3 3 ° N a n d 1 0 2 - 1 0 6 ° E in nor
thwestern Sichuan in 1976-1980. T h e s e specimens w e r e studied by Leng & Y i n ( 1 9 8 3 ) a n d ten s p e c i e s o f p h e - b o t o m i n e sandflies have b e e n identified including t w o n e w species - P. sicbuanensis and S. zhengjiani. At that time there w a s n o publication n o r personal c o m m u nication stating that these s p e c i m e n s h a d b e e n autho
rized by X i o n g as large type Pc which can not b e legally studied, n a m e d or r e n a m e d by others.
COMPARABILITY AND RELIABILITY
Following the updated m e t h o d o f p h l e b o t o m i n e t a x o nomy, P. sicbuanensis w a s carefully described and n a m e d following the International C o d e o f Zoological Nomenclature (Leng & Yin, 1 9 8 3 ) . T h e m o r p h o m e t r y and morphostructural c o m p a r i s o n o f twelve male cha
racters a n d six f e m a l e c h a r a c t e r s for four s p e c i e s recorded in Chinese Acilerius has s h o w n o b v i o u s spe
cific differences b e t w e e n Pc and Ps. ( T a b l e s I, III) In the papers o f ISS, there w e r e lack o f morphological descriptions and standards for supporting the diffe
rentiation b e t w e e n Ps and Pc. This will undoubtedly
m a k e o b s c u r e their s p e c i e s identification b e t w e e n Pc and Ps. Furthermore, many contradictory facts and data have b e e n given in the SEM studies carried out b y investigators from Shanxi (Zhao, Li, Q i a o & Yin, 1 9 9 7 ) . T h e y published four SEM imagines o f Pc female pha
ryngeal armatures which w e r e re-used and re-illustrated differently as to t h e site o f collection a n d the species naming without explanation b y Zhao, Q i a o , Yin, Y i n
& Li ( 1 9 9 8 ) , o f w h o m three w e r e the 1997 writers.
T h e o b s c u r e ness o f ISS works is also d e n o t e d in the magnification labeled in SEM imagines a n d their ratios to real size o f Pc a n d Ps. In the s p e c i m e n s collected from Nanping, northwestern Schuan (Zhao, Li, Q i a o &
Yin, 1 9 9 7 ) , the large type Pc ( P S ) in photo 1 ( 2 , 4 ( ) 0 x ) is almost the size o f small type Pc (Pc) in p h o t o 2 ( l , 6 0 0 x ) , w h i c h m e a n s that the real size ratio is about large type: small type = 2:3, Ps<Pc. T h e real size o f t h e s e t w o flies c a l c u l a t e d b y f e m a l e wing length ( T a b l e III) is Ps ( 2 7 5 0 ) : Pc ( 2 3 2 0 ) = 1.19:1.00, Ps>Pc.
T h e a b o v e confusions are clear just with a glance to T a b l e IV. It is impossible to evaluate the accuracy o f ISS's work and it would b e strange to treat Ps as a large lype o f Pc u p o n such o b s c u r e bases.
MALE ASCOID FORMULA
X i o n g et al. ( 1 9 9 0 ) have reported that they dissected 53 males from 1,500 m altitude a n d six males a b o v e 2,000 m altitude (for observing ascoid dissection is not n e e d e d , it is hard to understand w h y they have dis
sected the flies to observe ascoid)), and discovered that the ascoid formula o f large type Pc (Ps) has t w o pat
terns, 2 / 3 - 1 5 and 2 / 3 - 8 , 1 / 9 - 1 5 . But o n the basis o f e x a m i n i n g h u n d r e d s o f s p e c i m e n s from S i c h u a n , Yunnan and Tibet the authors claim that the ascoid for
mula o f male Ps is constantly 2 / 3 - 8 , 1 / 9 - 1 5 , in o n e pattern. In the view o f ascoid formula patterns among the m e m b e r s o f s u b g e n u s Adlerius over the world as s h o w n in T a b l e V, it is clear that they are different spe
cies with their o w n ascoid formula pattern. According to the o b s c u r e original s p e c i m e n s from a parapatric z o n e o b s e r v e d b y X i o n g et al. it is obvious that their wrong conclusion has c o m e from mixed original mate
rials o f Ps and Pc observed.
Ascoid formula Species
2 / 3 - 1 5 cbinensis, simici
2 / 3 - 5 , 1/6-15 angustus, balcanicus, comatus, hyreniae.
salangensis, zulfagarensis 2/3-7, 1/8-15 davidi, hindustanicus
2 / 3 - 8 . 1/9-15 ambitus brevis. fengi. halpensis. kabulen- sis, lotigiductits. ntjiesler. sicbuanensis, tu
rati ic us
Table V. - Ascoid formula of male subgenus Adlerius phlebotomine sandflies.
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L E N G Y A N J I A & Z H A N G L I N G M I N
Neglecting most papers meticulously w o r k e d under the light o f updated t a x o n o m y o n this item and published in world literature including Leng & colleagues studies, and hiding the characters o f identification described b y these investigators, ISS repeatedly sophisticated that Leng has only taken the difference o f male ascoid for
mula to name a normal (?) Pc as Ps (Xiong et al, 1990;
Z h a o et al, 1997; 1 9 9 8 ) . It is really a kind o f distort quotations c o m m o n l y used w h i c h would not e v e n alter facts. Scientific c o n c l u s i o n can only b e m a d e o n the basis o f facts and scientific data instead o f a hasty a n n o u n c e m e n t .
Z O O G E O G R A P H I C A L D I S T R I B U T I O N
It s e e m s that the C h i n e s e Adlerius p h l e b o t o m i n e s may b e divided into t w o groups according tomorphological approximation and geographical distribution. High Alti
tude Group: P.fengi, and P. sichuanensis. Low Altitude G r o u p : P. chinensis and P. longiductus stands in the middle ( T a b l e V I ) .
Species <5 AT o* A4 5 A4 9 CSD
P. chinensis 2 / 3 - 1 5 < 1 0 0 (34) < 1 0 0 (51) + P. longiductus 2 / 3 - 8 , 1/9-•15 < 1 0 0 (56) < 1 0 0 (66) + P. sichuanensis 2 / 3 - 8 , 1/9-•15 > 1 0 0 ( 2 0 3 ) > 100(148) - P. fengi 2 / 3 - 8 . 1/9--15 > 100 (257) > 100(178) - AF = ascoid formula, CSD = common spermathecal duct.
Table VI. - Some morphological differences in Chinese Adlerius.
T A X O N O M I C P R I N C I P L E A N D R U L E
Every c o n t e m p o r a r y p h l e b o t o m i n e taxonomist k n o w s that the m o r p h o l o g y o f female pharyngeal armature and s p e r m a t h e c a are s u b g e n u s characters for Adlerius, which are usually similar in m o r p h o l o g y with uncer
tain minute difference b e t w e e n s p e c i e s within the sub
g e n u s and are usually not used for identifying species.
Male characters are mainly used for identifying s p e c i e s in Adlerius, and o f course, in zoological taxonomy, dif
ferent m o r p h o l o g i c a l characters are used for identifi
cation in different level o f taxa and for different objects.
Xiong, Jin, Hong et al. ( 1 9 9 0 ) denied the classification system o f T h e o d o r ( 1 9 5 8 ) and Artemiev ( 1 9 8 0 ) o n sub
genus Adlerius without dealing with w h i c h characters should b e n e e d e d for this purpose; they affirmed that the main differential characters for Pc to other m e m bers in subgenus Adlerius axe 1. spermatheca, 2. female phatyngeal armature and 3- male genitalia especially the ratio o f a e d e a g u s tubercle distance to a e d e a g u s length, and said that these useful characters are not different between Pc and Ps. Updated phlebotomine taxonomists should b e s h o c k e d by the a b o v e unusual words.
Many i n c o m p l e t e and over-simplified description and inaccurate or inadequate c o m p a r i s o n for establishing
n e w p h l e b o t o m i n e s p e c i e s have also b e e n d o n e by s o m e investigators causing m u c h c h a o s . A n e w taxon has to b e created with a d e q u a t e descriptions, as sti
pulated in the International Code o f Zoological Nomen
clature: "to b e available every n e w scientific name...
must b e a c c o m p a n i e d by a description o f definition that states in words characters that are purposed to dif
ferentiate the taxon..." [ICZN ( 1 9 8 5 ) Article 13(01. For the brightness, the International C o d e had lead to the 1st International Symposium for P h l e b o t o m i n e Sand
flies (ISOPS I ) , 1 9 9 1 , R o m e w h e r e a standardization w a s finalized for describing p h l e b o t o m i n e s and the Chinese version has b e e n published in China (Leng, 1 9 9 1 ) . Unfortunately ISS n e g l e c t e d the principle and rule for t a x o n o m y and the standard o f ISOPS, they have laid undue emphasis o n the importance o f female pharyngeal armature and s p e r m a t h e c a for identifying p h l e b o t o m i n e s p e c i e s in c o m m o n . Zhao, Q i a o et al.
( 1 9 9 8 ) q u o t e d such idea o f X i o n g , J i n et al. ( 1 9 8 7 ) :
"Female pharyngeal armature is o n e o f most important morphological a c c o r d a n c e for identifying a s p e c i e s in subgenus Larroussius. A s p e c i e s can b e correctly iden
tified if c o m b i n e d with the m o r p h o l o g y o f sperma
theca". T h e n this idea w a s grafted o n Pc and Ps and a c c o r d i n g to the similarity o f pharyngeal armature u n d e r SEM imagines, Zhao, Q i a o et al. ( 1 9 9 7 ) repead- tedly c o n c l u d e d that Ps is nothing m o r e than a large type Pc. But the truth is that, under an i n c o n c e i v a b l e idea, SEM imagines c o u l d not prevent them to avoid inevitable mistakes.
It is strange, furthermore, that Hsiung ( X i o n g ) , G u a n
& J i n from Shanghai ( 1 9 8 1 ) simply c o m p a r e d t w o p h l e b o t o m i n e s sandflies in different s u b g e n e r a , S. sin- toni Pringle, 1 9 5 3 o f s u b g e n u s Sergentomyia s. str.
Franca & Parrot, 1 9 2 0 and a n e w l y found fly o f sub
g e n u s Parrotomyia T h e o d o r , 1 9 5 8 from southern Xin
jiang, instead o f c o m p a r i n g m e m b e r s within the s a m e s u b g e n u s , to create the latter as a n e w taxon, "Ser
gentomyia turfanensis sp. nov." Sergentomyia tur- fanensis is c o n s i d e r e d b e i n g S. (Parrotomyia)palesta- nensis Adler & T h e o d o r , 1 9 2 4 in m o r p h o l o g y and geographical distribution (Leng, 1 9 9 7 ) . D i s o b e y i n g the well-established t a x o n o m i c rules and law m a d e the p r o p o s e d n a m e invalid.
G e n e r a l rule and principle for t a x o n o m y h a v e b e e n e s t a b l i s h e d b y g e n e r a t i o n s o f t a x o n o m i s t s ( Z h e n , 1 9 8 7 ) . Present t a x o n o m i c system for p h l e b o t o m i n e sandflies has b e e n mainly formed within this century and is constantly improving. At present n e w t e c h niques of molecular b i o l o g y are often used for unders
tanding evolutionary t e n d e n c y o f a population and for p r o b i n g sibling s p e c i e s . G e n e r a l principle and rule, k n o w l e d g e a n d m e t h o d o f z o o l o g i c a l t a x o n o m y should always b e strictly followed. It has b e e n p r o v e d that the c o n t e m p o r a r y system for p h l e b o t o m i n e t a x o -
8 Mise au point Parasite, 2001, 8, 3-9
CHINESE PHLEBOTOMINE SANDFLIES
n o m y is effective a n d reflective t o natural history a n d must n o t b e n e g l e c t e d .
CONCLUSION
A
ccording t o t h e a b o v e m e n t i o n e d facts, it is taxonomically clear that P. sichuanensis is an isolate s p e c i e s instead o f a large type o f P. chi- nensis.REFERENCES
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Reçu le 31 août 1999 Accepté le 21 septembre 2000
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