IHB OpenIR  > 鱼类生物学及渔业生物技术研究中心  > 期刊论文
Three new Diplozoidae mitogenomes expose unusual compositional biases within the Monogenea class: implications for phylogenetic studies
Zhang, Dong1,2,3; Zou, Hong1,2; Wu, Shan G.1,2; Li, Ming1,2; Jakovlic, Ivan4; Zhang, Jin4; Chen, Rong4; Li, Wen X.1,2; Wang, Gui T.1,2
2018-09-03
Source PublicationBMC EVOLUTIONARY BIOLOGY
ISSN1471-2148
Volume18Issue:1Pages:17
Abstract

Background: As the topologies produced by previous molecular and morphological studies were contradictory and unstable (polytomy), evolutionary relationships within the Diplozoidae family and the Monogenea class (controversial relationships among the Discocotylinea, Microcotylinea and Gastrocotylinea suborders) remain unresolved. Complete mitogenomes carry a relatively large amount of information, sufficient to provide a much higher phylogenetic resolution than traditionally used morphological traits and/or single molecular markers. However, their implementation is hampered by the scarcity of available monogenean mitogenomes. Therefore, we sequenced and characterized mitogenomes belonging to three Diplozoidae family species, and conducted comparative genomic and phylogenomic analyses for the entire Monogenea class. Results: Taxonomic identification was inconclusive, so two of the species were identified merely to the genus level. The complete mitogenomes of Sindiplozoon sp. and Eudiplozoon sp. are 14,334 bp and 15,239 bp in size, respectively. Paradiplozoon opsariichthydis (15,385 bp) is incomplete: an approximately 2000 bp-long gap within a non-coding region could not be sequenced. Each genome contains the standard 36 genes (atp8 is missing). G+T content and the degree of GC- and AT-skews of these three mitogenome (and their individual elements) were higher than in other monogeneans. nad2, atp6 and nad6 were the most variable PCGs, whereas cox1, nad1 and cytb were the most conserved. Mitochondrial phylogenomics analysis, conducted using concatenated amino acid sequences of all PCGs, indicates that evolutionary relationships of the three genera are: (Eudiplozoon, (Paradiplozoon, Sindiplozoon)); and of the three suborders: (Discocotylinea, (Microcotylinea, Gastrocotylinea)). These intergeneric relationships were also supported by the skewness and principal component analyses. Conclusions: Our results show that nad2, atp6 and nad6 (fast-evolving) would be better candidates than cox1 (slow-evolving) for species identification and population genetics studies in Diplozoidae. Nucleotide bias and codon and amino acid usage patterns of the three diplozoid mitogenomes are more similar to cestodes and trematodes than to other monogenean flatworms. This unusual mutational bias was reflected in disproportionately long branches in the phylogram. Our study offsets the scarcity of molecular data for the subclass Polyopisthocotylea to some extent, and might provide important new insights into the evolutionary history of the three genera and three suborders.

KeywordPhylogenomics Mutational bias Molecular marker Long branch Amino acid usage Ancestral gene order reconstruction
DOI10.1186/s12862-018-1249-3
Funding OrganizationNational Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province
Indexed BySCI
Language英语
Funding ProjectNational Natural Science Foundation of China[31872604] ; National Natural Science Foundation of China[31572658] ; Earmarked Fund for China Agriculture Research System[CARS-45-15] ; Major Scientific and Technological Innovation Project of Hubei Province[2015ABA045]
WOS Research AreaEvolutionary Biology ; Genetics & Heredity
WOS SubjectEvolutionary Biology ; Genetics & Heredity
WOS IDWOS:000443376200004
WOS KeywordCOMPLETE MITOCHONDRIAL GENOME ; TRANSFER-RNA GENES ; PLATYHELMINTHES MONOGENEA ; GYRODACTYLUS-SALARIS ; MOLECULAR PHYLOGENY ; PARASITIC FLATWORMS ; RDNA SEQUENCES ; RAINBOW-TROUT ; DNA VARIATION ; EVOLUTION
PublisherBMC
Funding OrganizationNational Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; National Natural Science Foundation of China ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Earmarked Fund for China Agriculture Research System ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province ; Major Scientific and Technological Innovation Project of Hubei Province
Citation statistics
Cited Times:1[WOS]   [WOS Record]     [Related Records in WOS]
Document Type期刊论文
Identifierhttp://ir.ihb.ac.cn/handle/342005/29434
Collection鱼类生物学及渔业生物技术研究中心_期刊论文
Corresponding AuthorWang, Gui T.
Affiliation1.Chinese Acad Sci, Inst Hydrobiol, Minist Agr, Key Lab Aquaculture Dis Control, Wuhan 430072, Hubei, Peoples R China
2.Chinese Acad Sci, Inst Hydrobiol, State Key Lab Freshwater Ecol & Biotechnol, Wuhan 430072, Hubei, Peoples R China
3.Univ Chinese Acad Sci, Beijing, Peoples R China
4.Biotransduct Lab, Wuhan 430075, Hubei, Peoples R China
Recommended Citation
GB/T 7714
Zhang, Dong,Zou, Hong,Wu, Shan G.,et al. Three new Diplozoidae mitogenomes expose unusual compositional biases within the Monogenea class: implications for phylogenetic studies[J]. BMC EVOLUTIONARY BIOLOGY,2018,18(1):17.
APA Zhang, Dong.,Zou, Hong.,Wu, Shan G..,Li, Ming.,Jakovlic, Ivan.,...&Wang, Gui T..(2018).Three new Diplozoidae mitogenomes expose unusual compositional biases within the Monogenea class: implications for phylogenetic studies.BMC EVOLUTIONARY BIOLOGY,18(1),17.
MLA Zhang, Dong,et al."Three new Diplozoidae mitogenomes expose unusual compositional biases within the Monogenea class: implications for phylogenetic studies".BMC EVOLUTIONARY BIOLOGY 18.1(2018):17.
Files in This Item: Download All
File Name/Size DocType Version Access License
s12862-018-1249-3.pd(3434KB)期刊论文出版稿开放获取CC BY-NC-SAView Download
Related Services
Recommend this item
Bookmark
Usage statistics
Export to Endnote
Google Scholar
Similar articles in Google Scholar
[Zhang, Dong]'s Articles
[Zou, Hong]'s Articles
[Wu, Shan G.]'s Articles
Baidu academic
Similar articles in Baidu academic
[Zhang, Dong]'s Articles
[Zou, Hong]'s Articles
[Wu, Shan G.]'s Articles
Bing Scholar
Similar articles in Bing Scholar
[Zhang, Dong]'s Articles
[Zou, Hong]'s Articles
[Wu, Shan G.]'s Articles
Terms of Use
No data!
Social Bookmark/Share
File name: s12862-018-1249-3.pdf
Format: Adobe PDF
All comments (0)
No comment.
 

Items in the repository are protected by copyright, with all rights reserved, unless otherwise indicated.